Review of the genus Rhagovelia Mayr, 1865 (Insecta: Heteroptera: Veliidae) in the Palawan biogeographic region, the Philippines
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Zettel et al.: Rhagovelia (Veliidae) in the Palawan region, Philippines
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 68: 810–837
Date of publication: 9 November 2020
DOI: 10.26107/RBZ-2020-0092
http://zoobank.org/urn:lsid:zoobank.org:pub:6963D2F4-0E08-4455-AEFC-735A912DA735
Review of the genus Rhagovelia Mayr, 1865 (Insecta: Heteroptera:
Veliidae) in the Palawan biogeographic region, the Philippines
Herbert Zettel1*, Alice Laciny1 & Hendrik Freitag2
Abstract. This paper reports eight species of Rhagovelia Mayr, 1865, from the Palawan biogeographic region,
including three that are new to science. Rhagovelia minutissima Hungerford & Matsuda, 1961 (Busuanga Isl.);
R. estrella Zettel, 1994 (Central Palawan Isl.); R. palawanensis Zettel, 1994 (South and Central Palawan Isl.); R.
matillanoi, new species (North Palawan Isl.); and R. abbreviata, new species (North Palawan Isl.) are members
of the R. sarawakensis group, which is absent from other islands of the Philippine archipelago. Rhagovelia
kawakamii hoberlandti Hungerford & Matsuda, 1961 (Busuanga Isl., Palawan Isl., Balabac Isl., North Borneo) and
R. gapudi, new species (Busuanga Isl., Central and North Palawan Isl.) belong to the R. papuensis group (sensu
lato). Rhagovelia lansburyi Zettel, 1995 (Central Palawan Isl.) is an isolated species without clear affinity to other
species groups; thus, the Rhagovelia lansburyi species group is defined. The male genitalia and the macropterous
morph of R. minutissima are described for the first time. Numerous illustrations, maps, and keys to species and
species groups are provided.
Key words. Heteroptera, Veliidae, Rhagovelia, riffle bugs, new species, species groups, distribution, key, Palawan,
Busuanga, Balabac, Philippines
INTRODUCTION Biogeographic Regions. By geological history, it belongs
to the oldest landmasses of the country, the ‘Palawan
Riffle bugs of the genus Rhagovelia Mayr, 1865, are Microcontinental Block’, together with parts of the extant
frequently collected inhabitants of streams and rivers in islands of Mindoro, Romblon, and Panay. This block was
the tropics and subtropics. Approximately 400 species are originally part of the Eurasian continent, but separated about
described and many still undescribed species are known 32 million years ago by the opening of the South China Sea
from collections. This genus is well known for a high (e.g., Hall, 2002). There are indications that the block was
percentage of island endemics (e.g., Lundblad, 1936; never entirely submerged (e.g., Esselstyn et al., 2010) and
Nieser et al., 1997; Polhemus & Andersen, 2010, 2015) thus might have already carried initial faunal elements of East-
and therefore highly interesting for biogeographical and Asian continental origin. However, temporary land bridges
evolutionary studies. The Philippine species were revised in arising during periods of low sea levels in the Pleistocene
part by Zettel (1994b, 1995a, 1996). Several additions were have connected the islands of Greater Palawan with each
published later (Polhemus, 1996; Zettel, 1999, 2003a, 2007, other. Although the existence of a real land bridge was
2012, 2014; Zettel & Bongo, 2006). These studies treat the originally rejected (Voris, 2000), most likely this complex
five species hitherto known from the Palawan region. The must have been connected with the Sundaic region and the
newly studied material was mainly collected in the last two continent again (Esselstyn et al., 2010). It was proposed that
decades, primarily retrieved in connection with the AQUA these more recent geological events have shaped the faunal
Palawana Program and current surveys of the Biodiversity composition of Greater Palawan to a larger extent (Heaney,
Laboratory of the Ateneo de Manila University. 1985) and have caused more faunal similarity of Palawan with
Borneo than with the eastern Philippine Islands. The latter
The Palawan biogeographic region. Greater Palawan differs pattern was already recognised by Huxley (1868) for birds.
in its biogeographic history from all other Intra-Philippine While this was confirmed for some other vertebrates (e.g.,
Esselstyn et al., 2004), the observable patterns in insects are
more complex and depend on the dispersal abilities of the
taxa. Close affiliations of Palawan species with those from
Accepted by: Tran Anh Duc the eastern Philippines, Borneo, Eastern Asia, and Sulawesi
2 Zoological Department, Natural History Museum, Burgring 7, A-1010 Vienna,
1 nd are evident (e.g., Andersen, 1967; Holloway, 1982; Zettel,
Austria; Email: herbert.zettel@nhm-wien.ac.at (*corresponding author) 1994b, 1995b; Wang & McCafferty, 1995; Mey, 2001; Beck
Ateneo de Manila University, Biology Department, School of Science & Engineering,
2 et al., 2006; Braasch, 2011; Freitag et al., 2016).
Loyola Heights, Quezon City 1108, Philippines
© National University of Singapore
When considering the freshwater-inhabiting Hemiptera of the
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print) Palawan region at the species level, a very large portion of
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species appears to be endemic; for example, all species of and R. orientalis group are strongly dominating, while in
Helotrephidae (Zettel, 2003b), Aphelocheiridae (Polhemus Greater Palawan, the R. papuensis group is represented
DA & Polhemus JT, 1988), Notonectini (Notonectidae) by only two widely distributed species (R. kawakamii
(Nieser & Zettel, 1999), and Ptilomera Amyot & Serville, hoberlandti Hungerford & Matsuda, 1961; and R. gapudi,
1843 (Gerridae: Ptilomerinae) (Hungerford & Matsuda, new species), and the R. orientalis group is replaced by the
1965; Polhemus, 1998), as well as most species of Veliidae R. sarawakensis group. Field observations suggest that these
(although many of them are still undescribed; unpublished two groups occupy similar ecological niches. Rhagovelia
data). Apart from some widely distributed species (e.g., lansburyi from Central Palawan is a very peculiar species
in genera Limnogonus Stål, 1868, in Gerridae; Microvelia with unclear relationships (see above). Short diagnoses
Westwood, 1834, in Veliidae; Micronecta Kirkaldy, and a key to the three species groups are provided in the
1897, in Micronectidae; or Anisops Spinola, 1837, in taxonomic part.
Notonectidae: Anisopinae), Palawan shares only very few
species with Borneo (Aphelonecta philippina Zettel, 1995, in
Notonectidae: Aphelonectini; Rheumatogonus vantoli Chen MATERIAL AND METHODS
& Nieser, 2002, in Gerridae: Ptilomerinae), and even less
species with Mindoro, the most adjacent major Philippine Examined specimens were dry-mounted or preserved in
island. ethanol. Locality data for the present study and taxonomic
revisions published early (Zettel, 1994b, 1995a, 1996) were
Treatment of Rhagovelia species groups and species in collected from several museum and private collections over
the Philippines. Adults of almost all Rhagovelia species more than 25 years. Early research by the first author was
are dimorphic, with an apterous (wingless) and a—usually conducted within the framework of the Philippine Water Bug
rarer—macropterous (winged) morph. These two forms are Inventory Project (PWBIP) in cooperation with the University
structurally extremely different. Matsuda (1956) described of the Philippines, Los Baños (see Gapud & Zettel, 1999;
the new subgenus Neorhagovelia Matsuda, 1956, for those Zettel & Gapud, 2003). Later research by the third author
species which possess a short pronotum in the apterous and co-workers was conducted under the umbrella of the
morph. However, such splitting into two subgenera most AQUA Palawana Program (e.g., Freitag, 2004; Freitag &
likely does not reflect natural clades and was rejected by Yeo, 2004; Braasch & Freitag, 2008) in affiliation with the
most of the subsequent authors. Instead, a group concept Western Philippines University (formerly State Polytechnic
for the species of Rhagovelia in the Oriental, Malesian, College of Palawan). Later on, the project was moved to the
and Australian Regions was established by Polhemus & Ateneo de Manila University and is currently managed by
Polhemus (1988), and later modified in several minor details the Biodiversity Laboratory, for which the Palawan Region
by J.T. Polhemus (1990), D.A. Polhemus (1996), Nieser et is currently a major research focus (e.g., Freitag & Zettel,
al. (1997), and Zettel (2003a, 2006). A comparison of these 2013, 2014; Mey & Freitag, 2019, 2020). Specimens from
species groups with those groups, which inhabit the New each site were shared between participating institutions.
World, Africa, and Madagascar, is still lacking.
Repositories of specimens from the Palawan region:
For the Philippines, a key to the species groups was presented ADMU Ateneo de Manila University, Biodiversity
by Zettel (1994b). Later, D.A. Polhemus (1996) discussed Laboratory, School of Science & Engineering,
and confirmed the inclusion of R. werneri Hungerford & Quezon City, Philippines
Matsuda, 1961, in the R. novacaledonica group, but excluded BMNH The Natural History Museum, London, United
R. hoogstraali Hungerford & Matsuda, 1961, to form a Kingdom (formerly British Museum of Natural
group of its own. These two species are endemic to the History)
mountains of Mindanao Island. Zettel (1999) recorded the BPBM B.P. Bishop Museum, Honolulu, USA
R. borneensis group as new for the Philippines (R. fischeri FMNH Field Museum of Natural History, Chicago, USA
Zettel, 1999, from North Luzon). Zettel (2003a) proposed JTPC John T. Polhemus Collection (now part of the
that the R. caesia group (sensu Zettel, 1995a) is polyphyletic Smithsonian Institution, Washington D.C., USA)
and based on convergences (characters related to a special NCTN Nico Nieser Collection, Tiel, the Netherlands
mating behaviour with small “phoretic males”), and restricted NHMW Natural History Museum Vienna, Austria
it in a narrow sense to those species inhabiting New Guinea. NMHN Philippine National Museum of Natural History,
The Philippine species formerly included in that group were Manila, Philippines
then placed in new, separate clades, namely R. lansburyi NMPC National Museum, Prague, Czech Republic
Zettel, 1995, which is an isolated species close to the R. PCSD Palawan Council for Sustainable Development,
borneensis group; and R. phoretica Polhemus, 1996, and R. Puerto Princesa, Philippines
sallyae Zettel, 2003, which form another Philippine-endemic UPLB Museum of Natural History, University of the
group called the R. phoretica group (sensu Zettel, 2003a; not Philippines, Los Baños, Laguna, Philippines
sensu Polhemus & Polhemus, 2011) and might be related USC University of San Carlos, Cebu City, Philippines
to the R. orientalis group. ZCVA Zettel Collection, Vienna, Austria
ZMB Museum für Naturkunde, Berlin, Germany
In the Palawan Region, only three species groups have been ZMUC Zoological Museum, University of Copenhagen,
recorded so far. In the remaining Philippines, the R. papuensis Denmark
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Zettel et al.: Rhagovelia (Veliidae) in the Palawan region, Philippines
In the material sections and descriptions, dealate specimens 4. Forewing with 2 closed cells basally (Fig. 4C). Further characters
are included in the numbers of macropterous forms. see couplet 3................................................ R. lansburyi group
– Forewing with 3–4 closed cells reaching distal half of wing
Insects were examined with a Leica Wild M10 binocular (Figs. 3F, 6C)...........................................................................5
5. Forewing without whitish or yellowish streak (Fig. 3F).
microscope (max. 128× magnification). Parameres and
Abdomen entirely black (except in some species from
proctigers were studied under a Leica MZ12 compound Sulawesi)...............................................R. sarawakensis group
microscope. Drawings were made with the help of a camera – Forewing with whitish or yellowish streak (Fig. 5C). Abdomen
lucida attached to these microscopes. never entirely black.................................... R. papuensis group
6. Forewing broken at mid-length; 2 closed cells visible. Further
Stacked digital images were taken with a Leica DFC450 distinguishing characters see couplet 3...... R. lansburyi group
camera attached to a Leica Z16APO optics carrier, using – Forewing broken behind mid-length (Fig. 2C, D); 3–4 closed
Leica Application Suite V3.8. Images were stacked with cells visible............................................R. sarawakensis group
ZereneStacker 64-bit and processed with Adobe Photoshop
7.0. Key to the species of Rhagovelia in the Palawan region
Measurements were performed with a Nikon SMZ1500 1. Anterior margin of pronotum entirely yellowish; connexival
margin yellowish (Figs. 5A–D, 6A–C). Pronotum of apterous
binocular microscope at magnifications from 51.2× to 256×
morph much longer than eye length, with strongly convex hind
and are given in millimetres. They refer to the maximum margin, covering most of mesonotum (e.g., Fig. 5A). Forewing
length or width of the respective structure. Measurements blackish with light streak at base (e.g., Fig. 5C). Body length
on the trunk were done in dorsal view of specimens, except 2.8–3.9 mm. (III. R. papuensis group)....................................2
body lengths of winged specimens, which were measured – Pronotum black, with short orange or yellowish mark anteriorly;
in the lateral view excluding wings. connexival margin black (Figs. 1A–D, 2A–D, 3A–F, 4A–C).
Pronotum of apterous morph shorter than eye length, with
Terminology largely follows Polhemus & Polhemus (1988) straight hind margin; mesonotum largely exposed (e.g., Fig.
and subsequent authors (e.g., Zettel, 1996, 2012; Nieser et 1A). Forewing without light streak at base (e.g., Figs. 3F, 4C).
Body length 1.9–2.9 mm, except 3.3–3.5 mm in the females
al., 1997) publishing on taxonomy of Oriental Rhagovelia.
of R. lansburyi..........................................................................3
2. Anterior part of head black (not in freshly moulted specimens).
Maps were originally generated in DIVA-GIS 7.5 freeware Tergites 7 and 8 without shiny areas. Juga (pair of plates at
(https://www.diva-gis.org/about) and further edited with sides of labrum and rostrum base) with small black spinulae.
Corel Photopaint 10. Paramere of male distally broad, truncate (Fig. 7G). Apterous
female with connexival margins widely separated all over
length (Fig. 5B). Common and widespread in the region, also
TAXONOMY on Borneo.........................................R. kawakamii hoberlandti
– Anterior part of head (in frontal view) usually pale orange
to brown, rarely blackish. Juga without spinulae. Apterous
Key to species groups in the Palawan region
specimens with shiny areas on tergites 7 and 8 (more distinct in
female). Paramere of male distally slender, with small ventrally
Notes. This key is modified from Zettel (1994b). It is possible directed lobe (Fig. 7H). Connexival margin of apterous female
to identify species directly using the ‘Key to species’ provided strongly convergent near apex (Figs. 6B, 10G). Widespread in
within this publication (see below). the region.............................................. R. gapudi, new species
3. Stout species, strongly dimorphic; female much larger than
1. Apterous (wingless) specimen (e.g., Figs. 1A–D, 2A, B, 4A, male, in apterous morph 3.03–3.34 mm vs. 2.34–2.63 mm. Male
5A)............................................................................................2 with modified foreleg (Fig. 4A), protibia with distinct bend
– Macropterous (fully winged) specimen (e.g., Figs. 3F, 4C, at distal fourth. In male, dentition of metafemur reduced; in
5C)............................................................................................4 female, metafemur without dentition (Fig. 4B). (II. R. lansburyi
– Dealate (broken-winged) specimen (e.g., Figs. 2C, 3C, D; this group)......................................................................R. lansburyi
morph does not exist in R. papuensis group).........................6 – Usually slender; if stout, then much smaller. Body length
2. Pronotum long, covering mesonotum (almost) completely. 1.9–2.6 mm. Foreleg of males not modified, protibia more or
Metafemur of male strongly incrassate (Figs. 5A, 6A)............ less straight. In males, dentition of metafemur prominent (e.g.,
.................................................................... R. papuensis group Fig. 9A, C, D); in female reduced, but always present (e.g.,
– Pronotum short; mesonotum fully exposed (e.g., Fig. 1A–D). Fig. 9B, E). (II. R. sarawakensis group).................................4
Metafemur of male not or moderately incrassate (e.g., Figs. 4. Abdomen without spike-like setae. Sternites 3–6 (7) of male
1A, C, 4A)................................................................................3 with sharp median carina. Mesofemur of female with short
3. Male “phoretic”, much smaller than female, with foreleg of patch of long setae at depression on distal third....................5
male modified for grasping (incrassate profemur, strongly – Connexiva (and in apterous specimens also anterior tergites) with
bent protibia; Fig. 4A), but metafemur not enlarged and with numerous short and stout, spike-like setae (Fig. 10A). Sternites
reduced dentition (Fig. 4A). Female with mesopleural groove 3–5 of male without sharp median carina (a very obtuse carina
and medial humps on tergites 1–3.............. R. lansburyi group present in R. palawanensis). Mesofemur of female with brush
– Male and female subequal in size. Male with rich dentition on of long setae in distal half.......................................................6
metafemur (e.g., Figs. 1A, 3A). Female without mesopleural 5. Very stout species (Fig. 3C, E); in female, abdomen strongly
groove and medial humps on tergites 1–3................................ shortened, upcurved in apterous morph. In male, median carina
...............................................................R. sarawakensis group on sternite 7 reduced, weaker than on sternites 3–6. Species
from North Palawan.......................R. abbreviata, new species
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RAFFLES BULLETIN OF ZOOLOGY 2020
– Species of normal shape (Fig. 3A, B). Abdomen of female half of mesofemora of females. The other two species R.
not shortened. In male, median carina on sternite 7 prominent, estrella and R. abbreviata, new species, have some other
accentuated by paired impressions, stronger than on sternites important characters in common, e.g., carinate sternites and
2–6. Species from Central Palawan.......................... R. estrella paramere shape of the males, although the female of the
6. Mesocoxa, mesotrochanter, and metafemur entirely black (Fig.
latter species displays a very peculiar habitus. While the
1C, D). Species from South and Central Palawan....................
......................................................................... R. palawanensis species within each of these two clades show an allopatric
– Mesocoxa, base of mesotrochanter, and base of metafemur distribution in general, two species of the first and the second
yellow (Figs. 1A, B, 2A–D)....................................................7 clade, respectively, were occasionally collected in the same
7. Spike-like setae sparse, in apterous male scattered from streams, thus demonstrating that even within the same species
mesonotum to tergite 5, in apterous female until tergite 2, group there are sympatric, or even syntopic, species.
rarely forming a transverse row on tergite 1. Segment 8 of
male ventrally convex. Apterous female without distinct Rhagovelia minutissima Hungerford & Matsuda, 1961
constriction at base of sternite 7. Female without distinct hair (Figs. 1A, B, 7A, B, 11B)
tuft on connexival corner. Species from Busuanga Island.......
............................................................................R. minutissima
Rhagovelia (Neorhagovelia) minutissima Hungerford & Matsuda,
– Spike-like setae numerous, in apterous male from mesonotum
1961: 267–268.
to tergite 7, in apterous female until tergite 4, with distinct
Rhagovelia minutissima – Zettel, 1994b: 38–39; Chen et al.,
transverse row on tergite 1. Segment 8 of male ventrally flat.
2005: 439.
Apterous female with strong constriction at base of sternite 7.
Female with distinct tuft of black hairs at connexival corner
(Fig. 10A; indistinct in some macropterous individuals). Species Material reported by Zettel (1994b). Holotype (apterous
from North Palawan........................R. matillanoi, new species male, FMNH; genitalia missing) and allotype (apterous
female, FMNH) from Busuanga Island, Dimaniang,
Rhagovelia sarawakensis species group CNHM-Philippine Zoological Expedition. The former
barrio Dimaniang was not drawn in the map of Hoogstraal
Diagnosis. Body minute, length usually less than 3 mm; most (1951: 69), but geographic coordinates (12.0407148°N,
species slender. Body colour mainly black, pronotum with 120.1450728°E) were presented by Brown et al. (2018).
short transverse stripe anteriorly. Antennae and legs black,
often bright yellow at base. Metafemur of male moderately Additional material examined (ZCVA, UPLB, NHMW).
enlarged, but with rich dentition. Abdominal sternite 7 without 5 apterous males, 3 apterous females from Busuanga, 2 km
paired tubercles. Paramere of male usually long and slender, W Coron; 22 apterous males, 21 apterous females, 1
and with ventrally curved hook or lobe at apex, but very macropterous female, from Busuanga, Road km 13 WNW
short and reduced in Philippine taxa (Fig. 7A–E). Apterous Coron, Balulu Falls; 10 apterous males, 14 apterous females, 2
morph: Pronotum short, length shorter than eye length (e.g., macropterous males, 2 macropterous females, from Busuanga,
Figs. 1A–D, 2A, B), in few extra-Philippine species slightly 5 km NW Coron, Mabintangen River; 1 apterous male from
longer. Mesonotum fully or widely exposed. Macropterous Busuanga, W Borac.
morph: Wings uniformly dark. Forewing with 3 or 4 closed
cells that distally reach apical half of wing. Wings of common Diagnosis. Extremely small, body length of male 1.94–2.06
dealate specimens broken off in distal half, distal of cells mm (apterous), 2.03–2.39 mm (macropterous), female
(Figs. 2C, D, 3D). Abdominal carina reaching posterior 2.03–2.39 mm (apterous), 2.38–2.45 mm (macropterous).
margin of tergite 3 (rarely onto tergite 2 only). Mesocoxa, mesotrochanter, and base of metafemur yellow.
Short and stout, spike-like setae numerous on connexiva
Distribution. Species of this group are distributed from India and, in apterous morph, sparse and scattered on mesonotum
to Lombok, Borneo, and the southwestern Philippine Islands. and tergites (in male until tergite 5, in female until tergite
A single record from Sulawesi (Polhemus & Polhemus, 1988) 2; rarely with transverse row on tergite 1). Male: Sternites
appears doubtful (Nieser et al., 1997). John T. Polhemus without median carinae. Sternites 6 and 7 with weak paired
(1990) correctly included the widely distributed Rhagovelia impressions. Segment 8 ventrally convex. Paramere: Fig. 7A.
sumatrensis Lundblad, 1933, in this species group, which Apterous and macropterous females: Mesofemur with brush
covers almost the whole area of the group, except Borneo and of long setae in distal half. Apterous female: Abdomen not
the Philippines. Further species—many still undescribed— shortened, without distinct constriction at base of sternite
have restricted distributions on continental Southeast Asia, 7. Connexival corner (segment 7) without distinct hair tuft.
Sumatra, Borneo, and Greater Palawan.
Descriptive notes. Measurements: Apterous males (n =
Notes. The five Philippine species of this group, all restricted 10): Body length 1.94–2.06; maximum body width (at
to the Palawan Region, seem to form an isolated clade and metapleura) 0.78–0.84. Head width 0.56–0.59. Pronotum
are well distinguishable from the Bornean species by a length 0.16–0.19, width 0.61–0.66.
strong reduction of the parameres. Three species, Rhagovelia
minutissima, R. palawanensis, and R. matillanoi, new species, Apterous females (n = 10): Body length 2.03–2.39; maximum
share two very peculiar characteristics which are unknown body width (at metapleura) 0.86–0.98. Head width 0.59–0.66.
from other species of the R. sarawakensis group, i.e., spike- Pronotum length 0.16–0.20, width 0.63–0.72.
like setae on the abdomen and a brush of long setae in distal
813
Zettel et al.: Rhagovelia (Veliidae) in the Palawan region, Philippines
Fig. 1. Habitus of Rhagovelia species. A, R. minutissima, apterous male; B, R. minutissima, apterous female; C, R. palawanensis, apterous
male; D, R. palawanensis, apterous female.
Macropterous males, previously undescribed (n = 2): Body segment 6. Metafemur hardly slenderer when compared to
length 2.29–2.33; maximum body width (at pronotum) 1.04. apterous male. Metatibia straight. Tergite 7 shiny in middle.
Head width 0.61; pronotum length 0.95–0.98.
Macropterous females, previously undescribed (n = 3): Body
Colour: As in apterous male. Forewings black, without length 2.38–2.45; maximum body width (at pronotum) 1.08–
pale streak. 1.10. Head width 0.63–0.65. Pronotum length 0.95–1.03.
Pilosity: Pronotum behind humeri with relatively long Colour: As in macropterous male.
decumbent pilosity. Basal veins of forewing with moderately
long black pilosity. Pilosity: Setae on pronotum and forewings as in macropterous
male. Pilosity on tergites 1–7 absent (studied in a female
Structures: Pronotum large, with protruding humeri. Forewing with dissected wings), on tergite 8 reduced. Sternites 6–7 at
with 4 closed cells, in all specimens broken behind level of connexival margins with long, posteriorly directed setae, more
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RAFFLES BULLETIN OF ZOOLOGY 2020
prominently developed on sternite 7. Pilosity of gonocoxa males, 6 apterous females from southern Palawan, 12 km S
as in apterous morph with some inconspicuous moderately Brooke’s Point, Cabangaan, Tamlang River.
long pilosity dorsally and apically.
Additional material examined (JTPC, NHMW, UPLB,
Structures: Strongly deviating from apterous female as the PCSD). 1 apterous female from central Palawan, Puerto
abdomen is not constricted posteriorly. Pronotum large, Princesa, Iwahig, Balsahan River upstream of dam, 9°46′36″N,
with protruding humeri. Forewing as in male. Hindleg 118°39′55″E; 12 apterous males, 1 macropterous male, 11
as in apterous female. Abdomen with slightly and evenly apterous females from central Palawan, Montible, large river
convergent connexival margins. Tergite 7 anteriorly broader at km 29 on Napsan Road, 9°41′24.8″N, 118°37′26.3″E, 30 m
than tergite 8. In dissected specimen tergites 5–7 with large a.s.l.; 1 macropterous male from central Palawan, Montible,
shiny areas in middle. Tergite 8 almost entirely shiny, slightly Salakot Falls, 9°42′05″N, 118°31′18″E, 350 m a.s.l.; 1
shorter than wide. apterous male, 1 macropterous female from central Palawan,
Montible, stream at km 26 on Napsan Road, 9°42′09.3″N,
Morphological and comparative notes. Only the apterous 118°38′21.9″E, 50 m a.s.l.; 12 apterous males, 5 apterous
type specimens were previously described. Hungerford & females from central Palawan, Napsan, large stream at km
Matsuda (1961) noted that the holotype’s genitalia were lost 42 on Napsan Road, 9°41′28.5″N, 118°31′19.4″E, 270 m
during dissection. The genitalia depicted by Andersen (1967) a.s.l.; 13 apterous males, 23 apterous females from central
belong to a male of R. palawanensis (see Zettel, 1994b). Palawan, Taritien River at Trident Mine, 7 km NW of Narra;
Therefore, they remained hitherto undescribed. However, 1 apterous female from central Palawan, Narra, 5 km W town
the paramere of R. minutissima, here illustrated for the first proper, Taritien River, 9°19′11″N, 118°22′35″E, 100 m a.s.l.;
time (Fig. 7A), is very similar to that of R. palawanensis. 8 apterous males, 1 macropterous male, 11 apterous females,
This is the only species of the R. sarawakensis group on 1 macropterous female from southern Palawan, Mainit
the island of Busuanga. For characters distinguishing this Falls, 18 km NW of Brooks Point, 50 m a.s.l.; 22 apterous
species from its closest relatives on Palawan Island, see the males, 14 apterous females from southern Palawan, Brooke’s
key. Macropterous specimens are very rare; of 91 specimens Point, Sabsaban Falls, 8°53′20″N, 117°50′38″E, 100 m a.s.l.
studied only 5 (5.5 %) were macropterous (including 3
dealate specimens). Diagnosis. Very small, body length of male 2.02–2.33 mm
(apterous), 2.13–2.42 mm (macropterous); female 2.22–2.56
Habitats. Rhagovelia minutissima is known from several mm (apterous), 2.41–2.47 mm (macropterous). Mesocoxa,
small, moderately fast-flowing, permanent creeks. This mesotrochanter, and entire metafemur black. Short and stout,
species can probably inhabit a similarly wide spectrum of spike-like setae numerous on connexiva and, in apterous
habitats as R. palawanensis (see below). morph, also on mesonotum and tergites (in male until tergite
7, in female until tergite 4; with transverse row on tergite
Distribution. Busuanga Island (Fig. 11B). 1). Male: Sternites without or with obtuse median carinae.
Sternites 6 and 7 with weak or shallow paired impressions.
Rhagovelia palawanensis Zettel, 1994 Segment 8 ventrally flat. Paramere: Fig. 7B. Apterous and
(Figs. 1C, D, 7B, 11B, 12A) macropterous females: Mesofemur with brush of long setae
in distal half. Apterous female: Abdomen not shortened, with
Rhagovelia minutissima (misidentifications; nec minutissima strong constriction at base of sternite 7. Posterior edge of
Hungerford & Matsuda, 1961) – Andersen, 1967: 274, 276; laterotergite 5 without medioposteriorly directed long setae.
Polhemus & Polhemus, 1988: 165. Connexival corner (segment 7) with distinct tuft of black
Rhagovelia palawanenis Zettel, 1994b: 40–44; Chen et al., setae. Tergites 7 and 8 with long pilosity (usually dense).
2005: 439. Gonocoxa with prominent, long pilosity at dorsal edge.
Material reported by Zettel (1994b). Holotype (apterous Description. A detailed description was provided by Zettel
male, NHMW) from central Palawan, 7 km N Narra, (1994b).
Estrella Falls. Paratypes (NHMW, FMNH, NCTN, UPLB,
ZMUC): 46 apterous males, 9 macropterous males, 36 Descriptive notes. Measurements: Apterous males (n =
apterous females, 2 macropterous females, collected with 10): Body length 2.02–2.33; maximum body width (at
the holotype; 52 apterous males, 53 apterous females from metapleura) 0.77–0.91. Head width 0.56–0.61. Pronotum
central Palawan, 9 km W Puerto Princesa, Iwahig, Balsahan length 0.16–0.19, width 0.59–0.70.
River; 3 apterous males, 3 apterous females from central
Palawan, 20 km WSW Puerto Princesa, Montible River; 2 Apterous females (n = 10): Body length 2.22–2.56; maximum
apterous males, 1 macropterous male, 6 apterous females body width (at metapleura) 0.88–1.06. Head width 0.58–0.64.
from central Palawan, 20 km W Puerto Princesa, Tacduan Pronotum length 0.16–0.19, width 0.64–0.72.
Area, Tacduan River; 2 apterous males, 1 apterous female
from southern Palawan, 10 km NE Quezon, Tumarbon Falls; Macropterous males (n = 10): Body length 2.13–2.42;
1 apterous male, 2 apterous females from southern Palawan, maximum body width (at pronotum) 0.95–1.03. Head width
Brooke’s Point, Makagwa, Noona Dan Expedition; 6 apterous 0.55–0.63. Pronotum length 0.97–1.08.
815
Zettel et al.: Rhagovelia (Veliidae) in the Palawan region, Philippines
Macropterous females (n = 3): Body length 2.41–2.47; females from central Palawan, Puerto Princesa, Cabayugan,
maximum body width (at pronotum) 1.03–1.06. Head width SSW Martarpi, Cabayugan River, 10°09′46″N, 118°49′29″E;
0.61–0.63. Pronotum length 1.08–1.09. 1 macropterous (delate) from central Palawan, Puerto
Princesa, Concepcion, Tarabanan River, ca. 6 km upstream
Variations. Differences between populations are chiefly Highway, 10°03′47″N, 119°00′53″E, 200 m a.s.l.
found in apterous females and may indicate local genetic
diversification. The following characters are intraspecifically Type locality. Northern Palawan, Roxas municipality,
variable: the grade of posterior constriction of connexival barangay Bagong Bayan, Ilian Falls, 10°25′40″N,
margins; the density of long pilosity on tergites 7 and 8; and 119°33′25″E, 40 m a.s.l.
the presence or absence of black pilosity on laterotergites
4 and 5. Diagnosis. Very small, body length of male 1.97–2.11 mm
(apterous), 2.13–2.28 mm (macropterous), female 2.27–2.38
Comparative notes. See keys and comparative notes of R. mm (apterous), 2.27–2.47 mm (macropterous). Mesocoxa
matillanoi, new species. (usually), mesotrochanter, metatrochanter, and base of
metafemur yellow; mesotrochanter apically with black ring.
Habitats. Rhagovelia palawanensis inhabits a wide range Short, stout, spike-like setae numerous on connexiva in
of small to large permanent streams with moderate surface apterous morph, also on mesonotum and tergites (in male
velocity and unpolluted water, with or without canopy (Fig. until tergite 7, in female until tergite 4; with transverse
12A). Specimens prefer littoral areas, often between emergent row on tergite 1). Male: Sternites without median carina.
rocks, but still in slow runs, avoiding pool bays with slack Sternites 6 and 7 with distinct paired impressions, so that
water. They seem to be insensitive to sunlight exposure. the medial zone appears elevated. Segment 8 ventrally flat.
Paramere: Fig. 7C. Apterous and macropterous females:
Distribution. Palawan Island (South and Central) (Fig. 11B). Mesofemur with brush of long setae in distal half. Apterous
female: Abdomen (Fig. 10A, B) not shortened, with strong
Rhagovelia matillanoi, new species constriction at base of sternite 7. Posterior edge of laterotergite
(Figs. 2A–D, 7C, 8A, 9A, B, 10A–C, 11B, 14A–C) 5 with few medioposteriorly directed setae. Connexival corner
(segment 7) with distinct tuft of black setae. Tergites 7 and 8
Rhagovelia cf. palawanensis – Freitag & Zettel, 2013: 65–66. with long pilosity. Gonocoxa with few setae at dorsal edge.
Etymology. The species is named for Joie Matillano, Description of apterous male. Measurements of holotype:
formerly faculty member and collaboration partner of the Body length 2.06; maximum body width (at metapleura) 0.84.
senior authors, in recognition of his outstanding contributions Head length 0.28, width 0.61; minimum eye distance 0.15.
to the survey and protection of the Palawan fauna. We Pronotum length 0.19, width 0.66. Lengths of antennomeres:
particularly appreciate his dedicated encouragement of I 0.53, II 0.24, III 0.36, IV 0.35. Lengths of leg segments:
students in fieldwork and biodiversity studies, such as in our profemur 0.63, protibia 0.66, protarsus 0.02 + 0.02 + 0.16,
joint workshop “Freshwater Invertebrates: their Taxonomy, mesofemur 1.03, mesotibia 0.80, mesotarsus 0.05 + 0.31
Diversity and Ecology” in 2007, which also yielded additional + 0.50, metafemur 0.78, metatibia 0.73, metatarsus 0.04 +
data and material for this paper. 0.05 + 0.19.
Type material. Holotype (apterous male; NMNH) labelled Measurements of paratypes (n = 10): Body length 1.97–2.11;
“PHIL: Palawan, Roxas, Bagong Bayan, creek downstream maximum body width (at metapleura) 0.80–0.88. Head width
of Ilian Falls, 10°25′40″N 119°33′25″E, ca. 50 m a.s.l., 0.56–0.61. Pronotum length 0.16–0.19, width 0.63–0.69.
17.XI.1995, leg. J. Mendoza (# 129)”. Paratypes (ADMU,
NHMW, NMNH, UPLB, PCSD): 56 apterous males, 6 Colour: Black. Pronotum with transverse orange mark in
macropterous males, 76 apterous females, 11 macropterous centre, laterally reaching level of medial eye margin. Basal
females collected from the same locality (partly labelled third to half of antennomere 1, distal corner of proepisternum,
“Bagong Bayan Falls”); 1 apterous male, 5 macropterous all coxae (mesocoxa in some specimens infuscated) and
males from northern Palawan, El Nido, Barutuan, road km trochanters (mesotrochanter with distal black ring), basal half
291.5, small lowland river below irrigation dam, ca. 11°18′N, of profemur, and basal quarter to third of metafemur yellow.
119°27′E; 1 apterous male from northern Palawan, El Nido, Rostrum yellowish to dark brown. Juga brown to blackish.
Pasadeña, Nagkalit-Kalit Falls, ca. 11°15′N, 119°26′E; 24
apterous males, 3 macropterous males, 42 apterous females, Pilosity: Sides of pronotum, propleuron, and mesopleuron
2 macropterous females from northern Palawan, 20 km N with long black setae. Thorax dorsally with short greyish
Taytay, Pularaquin, Canequi Falls, 10°56′59″N, 119°28′09″E, pilosity. Abdominal tergites in addition to short appressed
25 m a.s.l.; 15 apterous males, 1 macropterous male, 28 pilosity with slightly longer, subdecumbent, black setae.
apterous females from central Palawan, Puerto Princesa, Short spike-like setae numerous on connexival margins
Marofinas, Panaguman River, 10°15′09″N, 118°58′03″E; 1 and all tergites, scattered and individually varying in their
macropterous female from central Palawan, Puerto Princesa, presence on lateral parts of sternites 3–7, on hind margins
Cabayugan, Cabayugan River S of Martarpi, 10°09′47″N, of thoracic nota (laterally) and on thoracic pleura (dorsally),
118°50′37″E, 37 m a.s.l.; 14 apterous males, 14 apterous in most specimens rather numerous on metapleuron.
816
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Fig. 2. Habitus of Rhagovelia matillanoi, new species. A, apterous male; B, apterous female; C, macropterous dealate, male; D, macropterous
dealate, female.
Structures: Head slightly broader than anterior margin of consisting of ca. 1–6 short teeth or spinules. Metatibia stout,
pronotum. Juga flat, inclined, and relatively broad, without straight or weakly curved, with strong dentition at flexor side.
spinulae. Pronotum very short, along midline about 0.7 times
as long as head. Mesonotum along midline about 3.0 times Laterotergites almost horizontal. Tergite 7 about 2.0 times as
as long as pronotum. Proepisterna bearing numerous small long as tergite 6, and 1.0 times as long as broad. Abdominal
black spinulae, in some specimens lacking in posterolateral venter without median carina. Sternites 6 and 7 with shallow,
corner. but distinct paired impressions, medial zone between them
appearing elevated. Proctiger slender, lateral wings weakly
Profemur on extensor side with very shallow concavity. developed (Fig. 8A). Paramere short, U-shaped, apex hardly
Metacoxa and metatrochanter with few small black spinulae. protruded (Fig. 7C).
Hindleg (Fig. 9A) rather uniform; femora moderately
thickened, longest spine near mid-length; basal row consisting Description of apterous female. Measurements of paratypes
of ca. 8–13 small spinules (in some specimens the distal (n = 10): Body length 2.27–2.38; maximum body width (at
spinule slightly longer than others); posterodistal row metapleura) 0.97–1.05. Head width 0.61–0.66. Pronotum
consisting of long first tooth followed by 5–8 teeth of strongly length 0.17–0.19, width 0.69–0.73.
decreasing length; anterodistal row often inconspicuous,
817
Zettel et al.: Rhagovelia (Veliidae) in the Palawan region, Philippines
Colour: as in male. Description of macropterous female. Measurements of
paratypes (n = 10): Body length 2.27–2.47; maximum body
Pilosity: Sides of pronotum, propleuron, and mesopleuron width (at pronotum) 1.03–1.08. Head width 0.61–0.64.
with long black setae. Pro- and mesonotum with dense fine Pronotum length 0.97–1.06.
appressed pilosity. Mesofemur with strongly developed
brush of long setae ventrally over distal half. Metanotum and Colour: As in macropterous male.
tergite 1 only with few appressed hairs. Tergites 2–6 bare;
tergites 7–8 with dense, posteriorly directed pilosity, that on Pilosity: Setae on pronotum and forewings as in macropterous
tergite 8 longer. Laterotergites with fine pilosity laterally, on male. Pilosity on tergite 7 absent, on tergite 8 reduced.
segment 4 longer and denser; posterior corner of segment 5 Sternites 5–7 at connexival margins with long, posteriorly
with a few posteromedially directed setae; posterior corner directed setae, most prominently developed on sternite 7 (Fig.
of segment 7 with distinct tuft of posterodorsally directed 10C); tuft at connexival corner developed, but weaker than in
black setae (Fig. 10B). Gonocoxa with numerous long setae apterous morph. Pilosity of gonocoxa as in apterous morph.
ventrally, but at most with few setae at dorsal edge. Short
and stout, spike-like setae numerous on connexiva until Structures: Strongly deviating from apterous female as the
segment 5, scattered between pronotum and tergite 4, on abdomen is not constricted posteriorly. Pronotum large, with
thoracic pleura, and lateral parts of sternites 2–5. protruding humeri. Forewing as in male. Hindleg slightly
slenderer than in apterous female. Abdomen with slightly
Structures: Body slightly broader than in male; abdomen and evenly convergent connexival margins. Tergites 7 and
more elongated. Head distinctly broader than anterior margin 8 of similar size, each slightly shorter than wide.
of pronotum. Pronotum very short, along midline about
half of head length. Mesonotum along midline about 3.7 Comparative notes. This new species forms a complex
times as long as pronotum. Mesonotum bulging, posteriorly with R. minutissima from Busuanga and R. palawanensis
declivitous; hind margin slightly concave at middle. from southern and central Palawan. Leg colour is identical
to R. minutissima and structural differences are listed in key
Mesofemur moderately flattened in distal half. Metafemur couplet 6. Structurally, R. matillanoi, new species, appears
(Fig. 9B) slender, without basal and anterodistal rows of more similar to R. palawanensis, and reliable differences
teeth; posterodistal row consisting of one long tooth at mid- (besides leg colour) were only detected in apterous females:
length of femur followed by 2–5 shorter teeth of decreasing In R. palawanensis, the gonocoxa bears a thick brush of
length. Metatibia toothless, straight. setae on its dorsal edge, which is lacking in R. matillanoi,
new species. In R. matillanoi, new species, the posterior
In dorsal aspect (Fig. 10A) abdomen broad anteriorly and corner of the laterotergite bears a few thin posteromedially
strongly narrowed posteriorly, with strong constriction at base directed setae (absent in very few specimens but possibly
of segment 7. Connexival margins strongly convergent until broken off), which are absent in R. palawanensis.
segment 5, subparallel at segments 6 and 7. Laterotergites 2–4
strongly inclined on segments, 5–7 nearly vertical. Tergites Habitats. This species was found in rather undisturbed,
1 and 4 slightly, transversely elevated, tergites 2 and 3 flat shaded streams running through forests (Fig. 14A, C). Along
or slightly depressed; tergite 5 trapezoidal, with median the Tarabanan River, and especially the well-studied
blunt carina; tergite 6 developed as a narrow stripe; tergite Cabayugan River in the Puerto Princesa Subterranean River
7 about 1.5 times as long as tergite 6 and about 1.5 times National Park, R. matillanoi, new species, was restricted
as long as wide; tergite 8 as long as tergite 7 and slightly to the upper, well-shaded river courses (see Freitag, 2004:
broader. Connexival corner in lateral view blunt. Gonocoxa sites “CR1”, “CR2”) in forested areas with heterogeneous
plate-like. Proctiger small, knob-like. streambed morphology and moderately to fast-flowing water.
At site “CR1”, all except one specimen were retrieved by
Description of macropterous male. Measurements of drift nets installed in a fast-flowing creek section.
paratypes (n = 8): Body length 2.13–2.28; maximum body
width (at pronotum) 0.97–1.05. Head width 0.58–0.63. Distribution. Palawan Island (North) (Fig. 11B).
Pronotum length 0.97–1.03.
Rhagovelia estrella Zettel, 1994
Colour: As in apterous male. Forewings black, without (Figs. 3A, B, 7D, 11C, 12A, 13A)
pale streak.
Rhagovelia estrella Zettel, 1994b: 44–45; Chen et al., 2005: 438.
Pilosity: Hind margin of pronotum with some moderately
long, posteriorly directed black setae. Basal veins of forewing Material reported by Zettel (1994b). Holotype (apterous
with long black pilosity. male, NHMW) and paratypes (46 apterous males, 20 apterous
females, 1 macropterous male, NHMW, FMNH, NCTN,
Structures: Pronotum large, with protruding humeri. Forewing BPBM, UPLB) from Palawan, 7 km N Narra, Estrella Falls.
with 3 or 4 closed cells, in dealate specimens broken behind
level of segment 6. Metafemur slender when compared to Additional material examined (ADMU, JTPC, NHMW,
apterous male. Metatibia straight or weakly curved. In dealate UPLB). 1 apterous male from Palawan, Taritien River at
specimens tergite 7 visible and shiny in middle. Trident Mine, 7 km NW of Narra; 1 apterous male from
818
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Fig. 3. Habitus of Rhagovelia species. R. estrella: A, apterous male; B, apterous female. R. abbreviata, new species: C, apterous male;
D, macropterous dealate, male; E, apterous female; F, macropterous female.
819Zettel et al.: Rhagovelia (Veliidae) in the Palawan region, Philippines
central Palawan, Montible, at Napsan Road km 29, large males, 9 apterous females, 5 macropterous females collected
river, 9°41′25″N, 118°37′26″E, 30 m a.s.l.; 31 apterous males, from the same locality (partly labelled “Bagong Bayan
14 apterous females from central Palawan, Napsan, Napsan Falls”); 1 apterous male, 1 apterous female from northern
Road km 43, Salakot Waterfalls, 9°42′10″N, 118°31′17″E, Palawan, El Nido, New Ibahay, mountain creek in disturbed
310 m a.s.l. primary forest, ca. 150 m a.s.l., 11°13′06″N, 119°29′32″E; 5
apterous males, 3 apterous females from northern Palawan,
Diagnosis. Small, body length of male 2.28–2.47 mm 20 km N Taytay, Pularaquin, Canequi Falls, 10°56′59″N,
(apterous), 2.50 mm (macropterous), female 2.53–2.75 119°28′09″E, 25 m a.s.l.
mm (apterous). Mesocoxa, (base of) mesotrochanter, and
base of metafemur yellow. Body without spike-like setae. Type locality. Northern Palawan, Roxas municipality,
Male: Sternites 2–7 with sharp median carinae. Sternite 7 barangay Bagong Bayan, Ilian Falls, 10°25′40″N,
with large paired impressions accentuating median carina. 119°33′25″E, 40 m a.s.l.
Segment 8 ventrally convex. Paramere: Fig. 7D. Apterous
and macropterous females: Mesofemur without brush of Diagnosis. Small, body stout, length of male 2.00–2.13 mm
long setae in distal half. Apterous female: Abdomen not (apterous), 2.14–2.27 mm (macropterous), female 1.92–2.19
shortened, without distinct constriction at base of sternite mm (apterous), 2.31–2.41 mm (macropterous). Mesocoxa,
7. Connexival corner (segment 7) without distinct hair tuft. base of mesotrochanter, and base of metafemur yellow. Body
without spike-like setae. Male: Sternites 3–6 with sharp
Descriptive notes. Measurements: Apterous males (n = median carinae. Sternite 7 with indistinct paired impressions,
10): Body length 2.28–2.47; maximum body width (at a median carina only indicated. Segment 8 ventrally convex.
metapleura) 0.91–0.97. Head width 0.66–0.72. Pronotum Paramere short, U-shaped, but apex protruded (Fig. 7E).
length 0.19–0.22, width 0.72–0.78. Apterous and macropterous females: Mesofemur with a short
depression at ¾5 of length, at this position ventrally with a
Apterous females (n = 10): Body length 2.53–2.75; maximum tuft of long setae. Apterous female: Abdomen (Fig. 10D,
body width (at metapleura) 1.03–1.11. Head width 0.69–0.75. E) strongly shortened, shorter than wide. Connexiva bearing
Pronotum length 0.19–0.22, width 0.73–0.81. brushes of black setae at segments 5 and 7. Tergite 6 very
short; laterotergite 6 absent; sternite 6 laterally reduced. Apex
Macropterous male (n = 1): Body length 2.50; maximum of abdomen starting from segment 7 directed dorsally, in
body width (at pronotum) 1.06. Head width 0.67. Pronotum two females from Pularaquin more posterodorsally. Tergite
length 1.06. 8 with long black pilosity posteromedially.
Morphological and comparative notes. Only a single Description of apterous male. Measurements of holotype:
macropterous individual is currently known (Zettel, 1994b). Body length 2.00; maximum body width (at metapleura) 0.88.
The carinate sternites of the males suggest a close relationship Head length 0.30, width 0.63; minimum eye distance 0.17.
of R. estrella and R. abbreviata, new species, but the females Pronotum length 0.22, width 0.72. Lengths of antennomeres:
of these two species are strongly different. For differentiation, I 0.51, II 0.28, III 0.40, IV 0.32. Lengths of leg segments:
see the characters listed in the key. profemur 0.66, protibia 0.70, protarsus 0.02 + 0.02 + 0.16,
mesofemur 1.09, mesotibia 0.94, mesotarsus 0.07 + 0.38
Habitats. The habitat requirements appear to be similar to that + 0.38, metafemur 0.78, metatibia 0.84, metatarsus 0.04 +
of R. palawanensis, as the two species partly occurred in the 0.04 + 0.21.
same collecting sites, including moderately fast-flowing, large
unshaded streams (Fig. 12A) as well as their smaller, more Measurements of paratypes (n = 10): Body length 2.02–2.13;
torrential tributaries (Fig. 13A). When occurring together, maximum body width (at metapleura) 0.81–0.88. Head width
R. estrella was less abundant than its congener. No records 0.61–0.64. Pronotum length 0.19–0.23, width 0.67–0.70.
on the particular microhabitats are available.
Colour: Black. Pronotum with transverse orange mark in
Distribution. Palawan Island (Central) (Fig. 11C). centre, laterally reaching level of medial eye margin. Basal
half of antennomere 1, distal corner of proepisternum, all
Rhagovelia abbreviata, new species acetabula, coxae, and trochanters (mesotrochanter with
(Figs. 3C–F, 7E, 8B, 9C–E, 10D–F, 11C, 14A–C) distal black ring), basal half of profemur, and basal fifth to
third of metafemur yellow. Rostrum and juga yellowish to
Etymology. The species epithet means “shortened” and refers dark brown. Middle of sternite 7 and segment 8 ventrally
to the peculiar shape of the female’s abdomen. yellowish brown.
Type material. Holotype (apterous male; NMNH) labelled Pilosity: Sides of pronotum, propleuron, and mesopleuron
“PHIL: Palawan, Roxas, Bagong Bayan, creek downstream of with comparatively short setae. Thorax dorsally with short
Ilian Falls, 10°25′40″N 119°33′25″E, ca. 50 m, 17.XI.1995, greyish pilosity. Abdominal tergites in addition to short
leg. J. Mendoza (# 129)”. Paratypes (in ADMU, NHMW, appressed pilosity with slightly longer, subdecumbent black
UPLB, PNM, PCSD): 22 apterous males, 8 macropterous setae. Entire body without short spike-like setae.
820RAFFLES BULLETIN OF ZOOLOGY 2020
Structures: Head slightly broader than anterior margin of in dorsal aspect. Segments 7 and 8 directed straight dorsad.
pronotum. Juga flat, inclined, and relatively narrow, without Tergites 1–6 short, visible as narrow stripes; tergite 7 about 3
spinulae. Pronotum very short, along midline about 0.7 times times as wide as long; tergite 8 largest. Sternite 6 normal at
as long as head. Mesonotum along midline about 2.5 times ventral part, laterally strongly narrowed towards connexival
as long as pronotum. Proepisterna bearing numerous small margins, and there hidden between sternites 5 and 7. Sternite
black spinulae anteriorly and mesally. 7 large, slightly flattened at middle. Gonocoxa large, evenly
convex. Proctiger small, narrower than long.
Profemur on extensor side with shallow concavity. Meso-
and metacoxa and metatrochanter with numerous small black Description of macropterous male. Measurements of
spinulae. Hindleg polymorphic: strong legs with moderately paratypes (n = 8): Body length 2.14–2.27; maximum body
enlarged femora and strongly curved tibia; slender legs with width (at pronotum) 1.00–1.03. Head width 0.61–0.64.
rather slender femora and almost straight tibia. Stout femora Pronotum length 1.00–1.09.
suddenly narrowed at distal third, basal row with 2–7 small
spines, large basal tooth of posterodistal row at first third of Colour: As in apterous male. Forewings black, without
femur length followed by 3–5 shorter teeth. Slender femora pale streak.
more evenly narrowed distally, basal row with 3–6 small
spines, posterodistal row consisting of 3–6 teeth. Anterodistal Pilosity: Apex of pronotal lobe with some long, posteriorly
row with fewer and much shorter teeth than posterodistal directed black setae. Basal veins of forewing with long
row. Metatibia with variable dentition at flexor side. black pilosity.
Laterotergites horizontal. Tergite 7 about 2.0 times as long Structures: Pronotum large, with protruding humeri. Forewing
as tergite 6, and 0.7 times as long as broad. Abdominal with 3 or 4 closed cells, in dealate specimens broken at
venter with sharp median carina at sternites 3–6. Sternite level of segment 6. Metafemur slender when compared to
7 with indistinct paired impressions, a median carina only apterous male. Metatibia straight or weakly curved. In dealate
indicated. Segment 8 ventrally convex. Proctiger slender, specimens, tergite 7 visible, which is shiny in middle.
lateral wings weakly developed (Fig. 8B). Paramere short,
U-shaped, but apex slightly protruded (Fig. 7E). Description of macropterous female. Measurements of
paratypes (n = 5): Body length 2.31–2.41; maximum body
Description of apterous female. Measurements of paratypes width (at pronotum) 1.09–1.13. Head width 0.66–0.67.
(n = 10): Body length 1.92–2.19; maximum body width (at Pronotum length 1.13–1.16.
metapleura) 0.97–1.02. Head width 0.66–0.69. Pronotum
length 0.22–0.25, width 0.69–0.75. Colour: As in apterous female. Forewings black, without
pale streak.
Colour: As in male, except propleuron with yellow patch.
Pilosity: Setae on pronotum and forewings as in macropterous
Pilosity: Pronotum laterally with long black setae. Mesofemur male. Sternites 5–7 at connexival margins with long,
with normal pilosity, except for a ventral (anterior) patch of posteriorly directed setae, most prominently developed on
long setae in the depression. Mesonotum and tergites 1–7 sternite 7.
without pilosity; tergite 8 with stiff, posteriorly-directed black
setae. Laterotergites with fine pilosity laterally; connexiva Structures: Strongly deviating from apterous female by
bearing brushes of short black setae at segments 5 and 7. a moderately shortened abdomen. Pronotum large, with
Gonocoxae and proctiger with fine short pilosity only. protruding humeri. Forewing as in male. Hindleg slightly
slenderer than in apterous female. Abdomen “normal”,
Structures: Body slightly broader than in male; abdomen directed straight caudad. Sternite 7 about twice as long as
strongly abbreviated. Head distinctly broader than anterior sternite 6.
margin of pronotum. Pronotum short, along midline about
0.8 times as long as head. Mesonotum along midline about 3 Comparative notes. The apterous female of R. abbreviata,
times as long as pronotum. Mesonotum posteriorly depressed; new species, characterised by the shortened and posteriorly
hind margin straight. upright abdomen, is unique among all species in Southeast
Asia, but a similar abdominal morphology is observed in
Mesofemur distally flattened, at ¾5 of length with unusual some distantly related species from Madagascar (Polhemus &
short depression (as if squeezed by forceps). Metafemur Andersen, 2010). The structure of the female’s mesofemur,
slender, without basal row of teeth, posterodistal row which bears a distinct depression and tuft of setae, is also
consisting of 1–3 teeth. Anterodistal row long consisting exceptional and a valuable character for recognition of
of minute, between pilosity hardly discernible spinules. winged females. Characters of the male (carinate sternites,
Metatibia toothless, straight. paramere with slightly protruded apex) suggest a close
relationship with R. estrella. These two species differ from
Abdomen ovoid in dorsal aspect. Connexival margins convex, other Philippine species of the R. sarawakensis group by the
strongly convergent until segment 5, segment 6 not visible absence of black spike-like setae on the abdomen.
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