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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4394 (4): 559
–
566
http://www.mapress.com/j/zt/
Article
559
https://doi.org/10.11646/zootaxa.4394.4.6
http://zoobank.org/urn:lsid:zoobank.org:pub:83AA97F4-AC3F-41AD-A339-5A67A64AF656
Description and notes on natural history of a new species of Parosus Sharp, 1887
(Coleoptera, Staphylinidae, Oxytelinae) living in floral bracts of
Columnea medicinalis L. (Gesneriaceae)
MARGARITA M. LÓPEZ-GARCÍA
1,2
& OSCAR H. MARÍN-GÓMEZ
1,2
1
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia.
E-mail: mamlopezga@unal.edu.co, ohmaring@unal.edu.co
2
Instituto de Ecología A.C., Carretera Antigua a Coatepec 351, El Haya, 91070 Xalapa, Veracruz, Mexico
Abstract
A new species of the recently revised genus Parosus is described, P. amayae López-García & Marín-Gómez sp. nov., from
adult and larval specimens collected in bracts of Columnea medicinalis in the Natural Reserve Río Ñambí (Southwestern
Colombia). Observations on the interaction with the plant, subsocial behavior, and population density are presented and
discussed. Adults and larvae apparently live together and feed on eggs and larvae of flies that develop inside the decom-
posing fruits of C. medicinalis. The new species is illustrated by color habitus photos, as well as its L1 and L3 larvae, male
and female genitalia are depicted by line drawings.
Key words: Oxytelinae, new species, decomposing fruits, gesneriads, subsocial behavior, cloud forest, Colombia, larvae
Introduction
Staphylinidae is the largest animal family with more than 61,300 described species (Newton 2015) and is dominant
in a great variety of ecosystems, showing several ecological interactions (Thayer 2005). However, biological
information is lacking for most species because of scarce field data, taxonomic limitations, and a high number of
undescribed species. Although associations with plants have been little documented, there are cases of phytophagy
(Klimaszewski et al. 2010), pollination (Ervik et al. 1999), hunting in phytotelmata (Frank & Barrera 2010), and
perching on the underside of leaves (López-García & Méndez-Rojas 2014). Rove beetles found on plants seldom
are phytophagous, but frequently saprophagous or predators (Thayer 2005). Another interesting aspect of the rove
beetle biology is the subsocial behavior which has evolved several times in Aleocharinae (Ashe 1987), Oxyporinae
(Setsuda 1994), and Oxytelinae (Hinton 1944). The subsociality refers to post-ovipositional parental behavior that
promotes the survival of offspring, requiring reproduction confined to specific periods and places, and considerable
adult longevity (Tallamy & Wood 1986). The known subsocial species of Oxytelinae live on dung, moist sand, or
intertidal mud, but associations with plants have not been well documented.
Among the flat-bodied Oxytelinae, the Neotropical genus Parosus Sharp, 1887 can be differentiated by a
median deeply emarginated labrum and a median serration on the posterior margin of the seventh tergum (Herman
1970; Makranczy 2014). The genus was described by Sharp and for a long time included only three species from
Panama and the Lesser Antilles (Sharp 1887; Herman 1970), but Makranczy (2014) described 17 additional species
and expanded the known geographical distribution of Parosus from Costa Rica to Southern Brazil. The genus is
restricted to cloud forests and Atlantic forest and some species seem to live in foliage (Makranczy 2014), but
information about collection methods is only available for half of the species and there were no specific data on the
vegetation where they live. One third of the species was described based on very few specimens (three or less), and
larval stages were unknown (Makranczy 2014).
In Colombia, there are only two described species of Parosus, collected from the Sierra Nevada de Santa
Marta, and lacking bionomic information. Samplings of arthropods associated with the species of the genus
Columnea L. (Gesneriaceae) from the Natural Reserve Río Ñambí, allow finding adults and larvae of a new species
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of Parosus that evidently live in the floral bracts of Columnea medicinalis (Wiehler) L.E. Skog & L.P. Kvist. Here,
we describe P. amayae López-García & Marín-Gómez sp. nov., including observations on the association with the
plant, subsocial behavior, and population density.
Material and methods
Taxonomy. The taxonomic keys by Herman (1970) and Kasule (1966) were used to identification of the adults to
genus and of the larvae to subfamily, respectively. The techniques for preparation and illustration of male genitalia
follow Makranczy (2006). Structure terminology and measurements follow Makranczy (2014). Measurements are
given in mm and were taken from ten well-developed specimens using an ocular micrometer. HW = head width
with eyes; TW = head width at temples; PW = maximum width of pronotum; SW = approximate width of elytra at
shoulders; MW = approximate maximum width of elytra; AW = maximum width of abdomen; HL = head length
from front margin of clypeus to the beginning of neck; EL = eye length; FL = faceted eye length; TL = length of
temple; PL = length of pronotum in the middle-line; SL = length of elytra from shoulder; SC = length of elytra
from hind apex of scutellum; FB = forebody length (combined length of head, pronotum, and elytra); BL =
approximate body length.
The type material was deposited at Instituto de Ciencias Naturales, Universidad Nacional de Colombia,
Bogotá, Colombia (ICN-MHN), Field Museum of Natural History, Chicago, IL, USA (FMNH), Muséum d’histoire
naturelle, Geneva, Switzerland (MHNG), and Naturhistorisches Museum Wien, Vienna, Austria (NHMW).
Observations on natural history. The Natural Reserve Río Ñambí is in the municipality of Barbacoas,
Nariño, Colombia (1º18’N 78º05’W; Fig. 1). This reserve is located in the Chocó Biogeographic Region, an
important global biodiversity hotspot (Myers et al. 2000) due the high levels of diversity and endemism (Rangel-
Ch 2015). It preserves a continuous fragment of 1400 ha of premontane pluvial forest with an elevational gradient
from 1100–1900 m.a.s.l. The annual average values for rainfall and temperature are 7160 mm and 19.2ºC,
respectively (Salaman 2001). The climatic conditions are very humid and wet reflecting a particular forest
physiognomy (Fig. 2): a canopy of 20 to 25 m dominated by palms, a dense understory layer and high density of
terrestrial and arboreal epiphytes (Salaman 2001; Marín-Gómez & Amaya-Márquez 2015; Rangel-Ch 2015).
FIGURES 1–2. 1) Known localities for Columnea medicinalis (GBIF 2017), location of the Natural Reserve Río Ñambí
(yellow circle), 2) Forest physiognomy in the Natural Reserve Río Ñambí.
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Columnea medicinalis is an endemic species from well preserved tropical to montane forests from
Southwestern Colombia and Northwestern Ecuador between 80–2400 m.a.l.s (Kvist & Skog 1993; Fig. 1). This
Gesneriaceae is an hemiepiphytic herb up to 4 m tall, with strongly dorsiventral stems and red apical spots on the
leaves (Figs. 7–8). The inflorescences are disposed in axils with 2–5 flowers (Fig. 9). The bracts are large, densely
congested, and cover up to 75% of the flower length. Flowers are white or yellow, tubular, and subventricose, with
two purple lines on the corolla lobes (Kvist & Skog 1993). The fruits are globosous whitish berries that ripe and
decompose inside the bracts, and the seeds can germinate there. In the Río Ñambí reserve, this plant has a high
population density (22 ind/ha), shows a clustering spatial distribution, and is exclusively pollinated by the Tawny
bellied hummingbird, Phaethornis syrmatophorus (Marín-Gómez & Amaya-Márquez 2015).
In July 2013 and February 2014, we sampled 18 individuals of C. medicinalis along its distribution (1100–
1500 m.a.s.l.) in the reserve. We collected one to three branches per plant and stored them immediately in a plastic
bag. Then we checked each sample removing one bract at a time looking for arthropods. When we found rove
beetles we recorded their movements along the bracts and their behavior. We collected all the arthropods in ethanol
(96%) and larvae were put later in a mixture of glycerol and ethanol (1:1). Some buds and fruits stored in FAA
were posteriorly dissected in laboratory.
Results
Taxonomy
Parosus amayae López-García & Marín-Gómez, sp. nov.
(Figs. 3–6, 10–12)
Type material. Holotype. Colombia, Nariño, Barbacoas, Altaquer, Reserva Natural Río Ñambí, brácteas de
Columnea medicinalis, 26.vii.2013, Col: O.H. Marín-Gómez (1 male, ICN-MHN). Paratypes (10 males, 14
females). Colombia, Nariño, Barbacoas, Altaquer, Reserva Natural Río Ñambí, brácteas de Columnea medicinalis,
26.vii.2013, Col: O.H. Marín-Gómez (2 males, 2 females ICN-MHN). Colombia, Nariño, Barbacoas, Altaquer,
Reserva Natural Río Ñambí, Brácteas de Columnea medicinalis (Gesneriaceae), 29.vii.2013, leg. O.H. Marín-
Gómez (3 males, 3 females, ICN-MHN). Colombia, Nariño, Barbacoas, Altaquer, Reserva Natural Río Ñambí,
Brácteas de Columnea medicinalis (Gesneriaceae), 26.vii.2013, leg. O.H. Marín-Gómez (1 dissected female,
FMNH). Colombia, Nariño dept., Barbacoas, Correg. Altaquer, Reserva Natural Río Ñambí, La Paila, 1º17’12.0"N
78º04’19.4"W, 1400 m, En brácteas de Columnea medicinalis, 3.ii.2014, leg. O.H. Marín-Gómez (1 male, 3
females, NHMW). Colombia, Nariño, Barbacoas, Correg. Altaquer, Reserva Natural Río Ñambí, En brácteas de
Columnea medicinalis, 4.ii.2014, leg. O.H. Marín-Gómez (2 males, 2 females, ICN-MHN, 1 male, FMNH, 1 male,
3 females, MHNG).
Larvae. Two L3 larvae collected with the holotype are deposited in FMNH (also deposited here a possibly
non-conspecific species, L1 larva and L3 larva, sampled at the same event). Further larvae (an L1 and L3 larvae
illustrated in Figs. 7-8) have the data "Colombia, Nariño, Barbacoas, Correg. Altaquer, Reserva Natural Río
Ñambí, En brácteas de Columnea medicinalis, 4.ii.2014, leg. O.H. Marín-Gómez" and are in various conditions
and completeness (in FMNH). The larval specimens are similar to those of Paraploderus (Figs. 78–79 in
Makranczy, 2016) except the very distinctive dark "stretch marks" found in the abdominal intersegments (Peter M.
Hammond, pers. comm.) that are present in Parosus and this is a common feature with Ochthephilus, Thinodromus
and Carpelimus, but lacking in Paraploderus.
Diagnosis. Parosus amayae López-García & Marín-Gómez sp. nov. is similar to Parosus rossii Makranzcy,
2014 from Ecuador on its color and general habitus. However, the new species can be differentiated by the
completely dark brown to black abdomen, larger body size (4.42–5.50 mm), and spoon shaped apexes of the
parameres (Fig. 4). In P. rossii the base of the abdomen is lighter, the body size is shorter (2.73–3.72 mm), and the
apexes of the parameres are wider and straight.
Description. Habitus as in Fig. 3. Measurements (n=10): HW = 1.02 (0.94–1.11); TW = 1.08 (1.01–1.15); PW
= 0.96 (0.89–1.04); SW = 0.87 (0.78–0.93); MW = 1.04 (0.89–1.13); AW = 0.86 (0.70–0.91); HL = 0.87 (0.79–
0.93); EL = 0.17 (0.16–0.21); FL = 0.12 (0.10–0.17); Tl = 0.32 (0.30–0.36); PL = 0.63 (0.57–0.69); SL = 0.91
(0.85–0.94); SC = 0.86 (0.83–0.90); FB = 2.67 (2.53–3.0); BL = 4.95 (4.42–5.50). Body 'bicolored'. Head black
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(supra-antennal prominences only slightly lighter), pronotum strongly reddish brown, elytra and abdomen black to
dark brown. Legs, mouthparts, and antennae light reddish brown to orange. Dorsal surface with medium short and
medium dense pale setae, slightly longer and sparse on frons, abdomen with much longer and sparser setae.
Head slightly wider than long. Mid-antennal articles moderately elongated (antennomere 6 length:width =
0.096:0.065 mm). Clypeus trapezoidal, ratio of longitudinal distance of supraantennal prominence tip from
eyefront to the same from clypeal front = 0.48–0.50. Infraocular ridge conspicuous, ending in a short keel after the
posterior ocular edge. Temple straight strongly curved at the posterior 2/3. Eye strongly bulging. Clypeus and
supraantennal ridges very shiny, with a few small, scattered punctures only. Vertex dense and strongly punctate.
Clypeus slightly elevated, frontoclypeal groove inconspicuous, only as a borderline before the strongly punctured
area. Head with about 32–34 'longitudinal' puncture lines, punctation becoming very dispersed in the triangle of the
supraantennal prominences and the mid-vertex.
Pronotum wider than long, maximum pronotal width 1.91x base width. Base straight, apex medially curved
upwards, sides curved, anterior pronotal angles slightly sharp. Pronotal midline slightly elevated, shiny, and
glabrous. Each side of midline broadly depressed. Pronotal disc densely punctate, punctures slightly smaller than
those on head. Pronotal margins with four widely separated macrosetae on each side, one on the apical margin and
the remaining three on the lateral margins.
FIGURES 3–6. Parosus amayae López-García & Marín-Gómez sp. nov. 3) Male habitus, 4) Right paramere in lateral view, 5)
Aedeagus in frontal view, 6) Spermatheca. Scale bar=0.1 mm for Figs. 4–6.
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FIGURES 7–12. Columnea medicinalis: 7) Hemiepiphytic habitus, 8) Detail of a branch, 9) Detail of an inflorescence.
Parosus amayae López-García & Marín-Gómez sp. nov.: 10) L1 and L3 larvae, 11–12) Larva and adult on bracts of C.
medicinalis.
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Elytra slightly dilated posteriorly, with two small, elongate impressions behind scutellum. Elytral punctation
not umbilicate, as large as on pronotum, separated by about 2 puncture diameters. Hind margin of tergite VII with a
medially serrate fringe. Aedeagus as in Fig. 4–5. Spermatheca as in Fig. 6.
Etymology. Parosus amayae López-García & Marín-Gómez sp. nov. was named after professor Marisol
Amaya-Márquez (Universidad Nacional de Colombia) who has studied the taxonomy and ecology of the species of
Columnea for more than twenty years and has made important contributions for knowing the diversity of
Gesneriaceae in Colombia.
Distribution. The species is only known from the Natural Reserve Río Ñambí (Nariño, Colombia).
Key to the described species of Parosus from Colombia
(based on original descriptions by Makranczy 2014)
1. Body bicolored; pronotum strongly reddish brown, head, elytra, and abdomen black to dark brown . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parosus amayae López-García & Marín-Gómez sp. nov.
1’. Body unicolored; head, pronotum, elytra, and abdomen black to dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Body length 3.23–3.58 mm. Infraocular ridge strong and thickened anteriorly, but vanishing posteriorly. Antennomere 6 about
as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Parosus colombiensis Makranczy, 2014
2’. Body length 4.55–5.40 mm. Infraocular ridge strong and continuing behind the eye. Antennomere 6 moderately elongated . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parosus gigantulus Makranczy, 2014
Natural history
Eighty-three individuals (48 adults and 35 larvae) of P. amayae López-García & Marín-Gómez sp. nov. were found
in Columnea medicinalis. The new species was observed in all the sampled plants (18 individuals), and its mean
density was three adults and two larvae per branch (five individuals per branch). The estimated population density
of the species in the studied area was 650 individuals/ha. Frequently, two well-developed individuals were found
along with a much smaller one, and two or three larvae in each branch. However, there were a few cases where
only one well-developed individual was accompanied by two small individuals. For the larvae, three stages were
identified with the following mean values of body length: L1 (1.9 mm; Fig. 10), L2 (3.8 mm), and L3 or mature
(5.2 mm; Fig. 10).
Adults and larvae remained hidden among the floral bracts of C. medicinalis, but sometimes they were moving
outside of the bracts (Figs. 11–12, video on Supplementary Material). They were also found inside the fruits and
sometimes an adult was alongside a larva among eggs and freshly hatched larvae of Diptera (Chironomidae). They
were not observed feeding on plant structures, but an adult was observed with one fly larvae in its mandibles. Other
insects were also found in the floral bracts of the plant but in very low abundance and in a few plants. They were
Staphylinidae (Hypotelus sp., Philonthina, Tachyporinae, Aleocharinae), Nitidulidae, Cantharidae, Formicidae
(Pheidole and Ectatomma), Hemiptera, Dermaptera, Blattidae, and larvae of Lepidoptera.
Discussion
Parosus amayae López-García & Marín-Gómez sp. nov. is just the third species of the genus known in Colombia,
but further samplings of specific plant microhabitats in cloud forests will allow discovering many others in the next
years. This kind of specific sampling will also help to collect the larvae of other Oxytelinae as very few have been
described so far (e.g. Makranczy 2016) and almost nothing is known about their biology. The present study
supports the suggestion of Makranczy (2014) that the species of Parosus live on foliage and their strongly flattened
body allow inhabiting among narrow microhabitats such as floral bracts. There is evidently a strong association
between P. amayae López-García & Marín-Gómez sp. nov. and C. medicinalis, but wider collections through the
distribution of this gesneriad, including North Ecuador (Fig. 1), are needed to prove the co-occurrence of the beetle
and the plant.
Columnea medicinalis is the only species of the genera in which the fruits decompose inside the floral bracts
providing a particular microhabitat. These bracts cumulate organic material and abundant food for P. amayae
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López-García & Marín-Gómez sp. nov. Although most Oxytelinae are saprophagous that feed on decaying plants
and algae (Thayer 2005; Makranczy 2006), the predator behavior has been found in adults and larvae of two
subsocial Platystethus that live on cattle dung (Palomino & Dale 1989; Hu & Frank 1995). However, the range
food of these oxyteline is not clear, and it has been not possible to known if they are coprophagous, mycophagous,
predators (Hu & Frank 1995), or a combination of all. In the case of P. amayae López-García & Marín-Gómez sp.
nov., the decaying fruits of C. medicinalis could be also a food resource, but only gut content analyses would allow
us to confirm it.
The subsocial parental care seems to evolve in response to unusually favorable environments that provide food
resources, which are rich but also scattered and ephemeral (Wilson 1971; Tallamy & Wood 1986). The
decomposing fruits of C. medicinalis provide a locally distributed resource for P. amayae López-García & Marín-
Gómez sp. nov., whether being predator or saprophagous, but also many other generalist predators can attack the
adults and the larvae of this species. As a defense mechanism, all the Oxytelinae have abdominal glands that
release secretions when are disturbed (Dettner & Schwinger 1982), which can be used by the adults of P. amayae
López-García & Marín-Gómez sp. nov. to avoid predators and protect the larvae. This territorial behavior could
play an important role in the reduction of floral herbivory (florivory) of C. medicinalis, as only a few herbivore ants
were collected in all the sampled bracts. The effects of florivory on pollination, the potential defense rol of P.
amayae against herbivores and the interaction with C. medicinalis along its distribution deserves further research.
Acknowledgements
Alfred Newton (FMNH) confirmed the genus identification. György Makranczy (Hungarian Natural History
Museum, Budapest) donated a paratype of P. rossii to ICN, gave valuable help during the description of the species,
and kindly contributed with the line drawings. Harald Schillhammer (NHMW) took the adult and larva habitus
photos. Howard Frank and Juan David Carvajal gave some logistical support. Klaus Mehltreter and Renato Portela
Salomão (Instituto de Ecología, A.C.) contributed improving the manuscript. The Natural Reserve Río Ñambí,
Jardín Botánico de Bogotá José Celestino Mutis, Nellie D. Sleeth Scholarship (The Gesneriad Society INC), and
Universidad Nacional de Colombia gave logistical and financial support for the development of this work.
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