the journal of

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NOTES ON GEOGRAPHIC DISTRIBUTION Check List 14 (5): 721-730

https://doi.org/10.15560/14.5.721

New and noteworthy records of rodents (Mammalia, Rodentia, Cricetidae and Echimyidae) from Paraguay

Robert D. Owen!?, Humberto Sanchez’, Karina Atkinson’, Leigh McMahon‘, Colleen B. Jonsson?

1 Dept. of Biological Sciences, Texas Tech University, Lubbock, Texas, USA. 2 Centro para el Desarrollo de la Investigacion Cientifica, Asuncion, Paraguay. 3 Instituto de Investigacion Biolégica del Paraguay, Calle Del Escudo N° 1670 casi Avenida Brasilia, Asuncion, Paraguay. 4 Para La Tierra, Centro IDEAL, Pilar, Mariscal Estigarribia 321 c/ Tte. Capurro, Neembucu, Paraguay. 4 Department of Microbiology, Immunology and Biochemistry, University of Tennessee Health Science Center College of Medicine, Memphis, TN, USA.

Corresponding author: Robert D. Owen, rowen@tigo.com.py

Abstract

Paraguay, situated in south-central South America, lies at the nexus of several major biomes, and exhibits high spe- cies richness for a relatively small country. Nevertheless, that faunal diversity 1s poorly known, and new species for the country continue to come to light. This paper reports on 3 species of rodents, 1 new for the country and 2 known previously in Paraguay from | specimen. We emphasize the need for more and better training of new field and museum biologists, as well as for directors of permitting agencies and protected areas.

Keywords Delomys, Interior Atlantic Forest; Juliomys, Kannabateomys;, Mbaracayu.

Academic editor: Alexandra M. R. Bezerra | Received 12 June 2018 | Accepted 17 August 2018 | Published 7 September 2018

Citation: Owen RD, Sanchez H, Atkinson K, McMahon L, Jonsson CB (2018) New and noteworthy records of rodents (Mammalia, Rodentia, Cricetidae and Echimyidae) from Paraguay. Check List 14 (5): 721-730. https://doi.org/10.15560/14.5.721

Introduction et al. (2002) listed 156 species, but increased collecting

Paani ellandlocked ecounttymin aouthiccnwalksouth activity and scrutiny of previously collected specimens

America, sits at the confluence of several major South American biomes—Pantanal, Interior Atlantic Forest, Cerrado, and Humid and Dry Chaco (Olson et al. 2001). Lying between ca 19 and 28 ° S, it is traversed by the Tropic of Capricorn, and thus includes both tropical and subtropical regions. Falling between ca 54 and 63° W longitude, the country includes a southeast to northwest annual precipitation gradient of from 1800-400 mm. Not surprisingly the country harbors a high diversity of animal species, including mammals, with 181 native mammal species currently recognized (de la Sancha et ale 20197);

The number of recognized mammal species in Para- guay has risen markedly during the past 2 decades. Myers

has revealed that several species’ distributions thought to terminate in Brazil or Argentina, actually extend into Paraguay. Most of these new records for the country were didelphid marsupials (Voss et al. 2009, de la Sancha and D’Elia 2015), bats (Stevens et al. 2010, Owen et al. 2014), and rodents (D’ Elia et al. 2008, de la Sancha et al. 2009), but some have also been larger species (Smith et al. 2011, 2013).

Of the 181 Paraguayan mammal species, 34 (repre- senting 20 genera) pertain to the subfamily Sigmodontinae (family Cricetidae), a widespread and species-rich radia- tion in the Neotropics. Another 22 species, representing 9 families, bring the total to 56 recognized Paraguayan rodent species. Among these, at least 2 species are

Copyright Owen et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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recorded in Paraguay from only | specimen. This report adds 1 new genus and species to the list of Paraguayan rodents, and provides new specimen localities for 2 which are known from only | specimen.

Methods

A long-term project on small mammal and Hantavirus ecology and evolution by personnel from Texas Tech University and the University of Tennessee Health Sci- ences Center, Memphis, has involved multiyear sampling of small mammals in the Reserva Natural del Bosque Mbaracayu (RNBM), Canindeyt' Department, northeast- ern Paraguay. The area of the reserve is approximately 65,000 ha. Most of the vegetation, including the area where we sampled, is mature Upper Parana Atlantic For- est (UPAF), some of which was selectively logged in the past, but undisturbed for at least the last 35 years. Our sampling has included both collecting and mark-recap- ture sampling, and has been conducted in various areas within and around the reserve during the years 1995-96, 2001-08, and 2014-17. Our database includes approxi- mately 7,780 capture records from the RNBM, and has revealed a high diversity both of sigmodontine rodents and small didelphid marsupials (Barreto Caceres 2017).

Most recently (2015-17), 6 sampling grids were established to monitor small mammal population levels and community composition (and Hantavirus serop- revalence) on a seasonal basis, through 2 years. Each grid consisted of 12 x 12 = 144 trap stations, 10 m apart, and thus sampled approximately 1.44 ha of the forest. Each station included 2 standard Sherman™ traps (7.6 x 8.9 x 22.9 cm; Sherman Trap Company, Tallahassee, FL) placed about 1 m apart on the ground at the station point, and another trap placed 2—3 m above the ground on branches or in vines as near as possible to the station point as vegetation permitted. Each grid was run for 5 nights during 6 sampling sessions, for a total of 77,760 trap-nights. In addition, 16 pitfall traps were situated uniformly in a4 x 4= 16 pattern on 3 of the grids. These traps were opened for 3 consecutive nights in August, September, October, and December 2016, and January 2017 (16 traps, 3 grids, 3 nights, 5 sessions, for a total of 720 pitfall trap-nights).

In a separate research program, Para La Tierra (a research-oriented non-governmental organization created in 2010 and based in Paraguay), has involved approxi- mately 310 volunteers, interns, visiting scientists and staff in developing a zoological collection including both invertebrates and vertebrates. Most of their collecting has been in departamentos San Pedro (adjacent to Canindeyu) and Neembucu (southwestern Paraguay), and they also collect opportunistically when animals are encountered elsewhere.

Specimens reported in this article were collected under Permisos de Colecta Cientifica N°. 011/2014, 132/2015, 269/2016, and 141/2017. Animals which were collected alive were euthanized following the Animal

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Care and Use Committee guidelines of the American Society of Mammalogists for the use of wild mammals in research and education (Sikes et al. 2011, 2016). All ani- mal procedures were approved (Approval No. 14024-03) by the Texas Tech University Institutional Animal Care and Use Committee (IACUC), which follows the 8th Edition of the Guide for the Care and Use of Laboratory Animals (Guide), NRC 2011. The study did not involve endangered or protected species. Specimens which are deposited in the collection of R. Owen are housed there temporarily by request of the Museo Nacional de Histo- ria Natural del Paraguay, where they will be permanently deposited when the museum has the resources to house them.

One new sigmodontine species is reported here for Paraguay, and new specimens and localities are reported from 2 additional rodent species that were previously known from only 1 specimen from Paraguay (Fig. 1). For each of the 3 species, collecting data are provided, as well as a description of the new specimens, followed by information on reproductive status, habitat and associa- tions with other species. Nomenclature follows Patton et al. (2015) and authors therein.

Results New records.

Delomys dorsalis (Hensel, 1872). Paraguay, Canindeyu Department, Reserva Natural del Bosque Mbaracayu (23.9964° S, 055.3671° W), collected by R. Owen and I. Mora, 6 October 2003, 1 adult female, R. Owen col- lection, field number TK 121143. Paraguay, Canindeyu Department, Reserva Natural del Bosque Mbaracayu (24.1424° S, 055.3454° W), collected by R. Owen, H. Sanchez, and C. Jonsson, 27 August 2014, 1 adult male, R. Owen collection, field number TK 184693.

Juliomys cf. pictipes (Osgood, 1933). Paraguay, Canindeyu Department, Reserva Natural del Bosque Mbaracayu (24.1410° S, 055.3663° W), collected by R. Owen and H. Sanchez, 15 July 2016, one adult male, R. Owen collection, field number 170166. Paraguay, Canindeytu Department, Reserva Natural del Bosque Mbaracayu (24.1414° S, 055.3659° W), collected by R. Owen, H. Sanchez and C. Jonsson, 15 February 2017, 1 adult female, R. Owen collection, field number TK 186360.

Kannabateomys amblyonyx (Wagner, 1845). Para- guay, Itapua Department, 7.9 km southeast of Alto Vera (26.8111° S, 055.7201° W), collected by K. Atkinson and L. McMahon, 22 October 2017, 1 subadult male, Zoo- logical Collections of Para La Tierra, CZPLT 486.

Comments and identification. The adult female De/o- mys dorsalis (TK 121143), with vagina closed, was captured in a Sherman trap placed on the ground. The locality is in a transitional area between bamboo forest (Guadua angustifolia) and primary semideciduous for- est of medium height, along the edges of Arroyo Karapa

Owen etal. | Records of rodents from Paraguay

bad

54°0'0"W

58°0'O"'W 56°0'0"W o) o o N Bolivia Argentina : er : a a Upper Parana Atlantic Forests ae yy Eo] g =. o “ 2 Oo o So al Argentina A Legend Rivers Cerrado Humid Chaco 1:2,500,000 ; es LK Upper Parana Atlantic Forests ME Lop 0 25 50 100 54°0'O"W

56°0'O"W

58°0'O"'W Figure 1. Map of eastern Paraguay showing major biomes and localities of specimens reported in this paper. The map also indicates nearest previously reported localities for the same species. Localities are: circles, Juliomys pictipes, TK 170166, TK 186360 (new localities, closed circle)

and FACEN-CZ 014 (previous locality [de la Sancha et al. 2009], open circle); squares, Delomys dorsalis, TK 141143, TK 184693 (new localities,

closed squares) and FMNH 26818 (previous locality [Goncalves and Oliveira 2014], open square); stars, Kannabateomys amblyonyx, CZPLT 486 (new locality, closed star) and BMNH 1903.4.7.20 (previous locality [Thomas 1903], open star).

in the upper reaches of an escarpment known as the megacephalus, which the species strongly resembles Cordillera Mbaracayu. The specimen was prepared as a externally. Later examination of the cleaned skull and fluid-preserved body with skull extracted and cleaned, re-examination of the body revealed that it represented and was initially identified and cataloged as Hylaeamys a species not yet recorded from Paraguay, which was

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Figure 2. Dorsal and ventral views of the crania of 2 Delomys dorsalis and 1 Hylaeamys megacephalus. Left, TK 121143; center, TK 184693

(both D. dorsalis); right, TK 130631 (H. megacephalus). See Table 1 for measurements of the D. dorsalis skulls.

subsequently determined to be De/omys dorsalis.

The adult male D. dorsalis (TK 184693), with scro- tal testes (15 x 7 mm) was collected in a Sherman trap placed on the ground at a locality ca 16 km south of the first specimen. This was in a trapping transect in an area of the RNBM characterized as bamboo understory, which typically is a high forest with a dense understory pre- dominantly of Merostachys clausseni bamboo, 2—3 m in height (Naidoo and Hill, 2006). The animal was prepared as a Standard skin and skull specimen.

These 2 specimens are clearly distinguished from externally similar species (principally Hylaeamys mega- cephalus) by a number of skull characteristics, the most obvious being the much shorter palatine foramina found in Hylaeamys, and the distinctly longer rostral tube in Delomys (Fig. 2). The interorbital region in H. megaceph- alus is generally ridged, but 1s smooth in these 2 De/omys

specimens. Externally, the mystacial vibrissae of the Delomys extend beyond the bases of the ears, whereas in Hylaeamys the vibrissae do not reach the ears. The claw of hind-foot digit I in Delomys extends well beyond the first interphalangeal joint in digit I, but barely reaches it in Hylaeamys. Ventral pelage of our dry specimen of Delomys is markedly dark gray-based and white-tipped, whereas that of H. megacephalus from the same locality shows less contrast between base and tip. The darker mid- dorsal pelage often observed in De/omys is minimally but clearly visible in our dry specimen (Fig. 3). Based on both geographic proximity and external morphologic char- acters, we identify these specimens as D. dorsalis. The tail of the dry skin specimen is only weakly bicolored, and the dorsal pelage is generally dark-cinnamon, with a moderately well-defined darker mid-dorsal stripe. The nearest collecting locality for another species of Delomys

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= = i) —

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Figure 3. Dorsal and ventral views of study skins of Delomys dorsalis and Hylaeamys megacephalus. Left, TK 121143, Delomys dorsalis; right, TK 130621, Hylaeamys megacephalus. Note the faint dark mid-dorsal band on the Delomys.

(D. sublineatus) is approximately 690 km distant, near the Atlantic coast in Parana state, Brazil (Goncalves and Oliveira 2014). External and cranial measurements of these 2 specimens are provided in Table 1.

Other species collected in the same trap line as TK 121143 were Akodon montensis, Oligoryzomys nigripes, and Oligoryzomys sp. Species collected in the same sampling line as TK 184693 were Akodon montensis, Hylaeamys megacephalus, Oligoryzomys nigripes, and Sooretamys angouya.

These new specimens represent a distribution extension of about 285 km north-northwest from the nearest previ- ously known locality (Caraguatay, Misiones Province, Argentina—the only record from Argentina) (Gongalves and Oliveira 2014). More importantly, these are the first localities recorded from west of the Parana River, and are the first reported from the Paraguay River watershed.

The adult male Juliomys (TK 170166), with scrotal testes was collected in the dry season, and the adult female with vagina closed was collected in the wet season. Both of these animals were captured in Sherman traps set 2—3 m above the ground, on branches or lianas. The locali- ties of these 2 captures were approximately 60 m apart. These records are approximately 115 km northwest from the locality of the only other Paraguayan specimen, and also are the first records of the species in the Paraguay River watershed.

These animals were easily distinguished from other possible species (principally Oligoryzomys nigripes) by their reddish rumps and rostra, tails distinctly bicolored and shorter than those of Oligoryzomys, and especially the reddish-orange coloration of the dorsal pelage, and pronounced white ungual tufts, of their hind feet (Fig. 4). Based on geographical proximity we tentatively iden- tify these specimens as J. pictipes. The nearest locality recorded for another species of Juliomys (J. ossitenuis) is about 540 km away, in Parana state, Brazil (Christoff et al. 2016). Nevertheless, our 2 specimens appear to more closely concur with the description of J. ossitenuis (Pavan and Leite 2011), and direct comparisons with other speci- mens are warranted. Although we report these specimens as a noteworthy locality for J. pictipes, re-indentification as J. ossitenuis would add yet another species to the list of Paraguayan mammals. The 2 individuals reported here were prepared as standard skin and skull preparations. External and cranial measurements of these 2 specimens are provided in Table 1. Measurements of the other Para- guayan specimen (FACEN-CZ 014) were listed in de la Sancha et al. (2009). We were unable to compare the prepared skins of our specimens with that of the other Paraguayan specimen, as that skin cannot be located. Photographs of the skulls of the 3 Paraguayan specimens are shown in Fig. 5.

Both of these individuals were collected on the same

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Table 1. External and cranial measurements from 5 specimens of 3 species reported in this paper. External measurements in millimeters, and weight in grams, taken from specimen tag and/or collector's notes. Cranial measurements in millimeters taken following Tribe (1996) by R.

Owen, except for CZPLT 486, taken by L. McMahon.

Juliomys cf. pictipes

Characters

TK 170166 TK 186360 Total 191 190 Tail 97 93 Hind foot 20 18 Ear 15 14 Weight 25 19 ONL 24.41 25.64 PL 10.93 11.47 PPL 8.81 9.34 MRC 3.50 3.91 M1B 1.03 1.16 PBL 3.68 3.67 TFL 7.59 7.12 DL 6.01 6.20 IFL 4.00 4.88 IFB 1.79 2.00 PB1 2.01 2.23 PB3 2.35 2.85 MFB 1.44 1.66 BIT 1.71 1.77 BW 3.42 3.76 BL 5.34 5.66 BCB 11.25 11.82 SH 7.41 7.88 RH 4.64 5.09 RB 4.26 4.64 RL 7.80 7.85 NL 8.78 8.92 ZPL 1.83 2.06 IOB 4.12 4.09 ZB 13.05 13.77 GLM 12.95 12.58 MMR 4.08 3.70 DR 2.42 2.49

sampling grid, which is characterized as dense forest with abundant lianas, mostly of family Bignoniaceae, associ- ated with large trees. The ground slopes northeasterly with a maximum slope of 6°. Mature trees reach an average height of 15 m. The upper canopy reaches 24 m in height and several of the tree species are deciduous. Astronium fraxinifolium is the most characteristic species, followed by Ocotea puberula, Peltophorum dubium, Inga affinis, and Parapiptadenia rigida. The lower arboreal stratum is characterized by Guarea kunthiana, O. puberula, and Chrysophyllum gonocarpum. Other species that appear less frequently are Nectandra megapotamica, Cabralea canjerana, Cariniana estrellensis, and Rheedia brasil- iensis. On average these species reach 8 m and grow associated with various species of lianas. The most frequent species of the shrub layer, which measures an average of 3.1 m in height, are Guarea kunthiana, Soro- cea bonplandii, and Rheedia brasiliensis. It is common to find regenerations of Nectandra megapotamica of no more than 2 m and dense groups of Piper hispidum some- times associated with P. amalago. In the clearings left by

Delomys dorsalis Kannabateomys amblyonyx TK 121143 TK 184693 CZPLT 486 255 262 328 126 132 194 30 28 40 20 21 18 42 51 97 32.64 32.84 44.35 14.47 14.25 18.49 11.61 11.98 NA 4.57 4.40 8.29 1.33 1.35 4.53 5.18 4.79 9.69 10.05 9.65 12.88 8.65 8.69 9.06 6.09 6.49 8.25 2.30 2.95 2.34 3.04 3.05 1.42 3.37 3.47 1.89 2,25 2.48 NA 2.17 1.73 2.65 4.63 3.65 oes) 4.52 4.04 NA 12.43 13.34 19.25 8.48 8.70 13.77 5.67 5.57 9.00 5.80 5.78 6.77 12.15 12.12 13.25 13.66 13.14 12.39 2.86 275 6.60 Sl? 4.97 11.85 1537 15.54 22.08 14.62 14.95 23.88 4.86 4.74 9.49 2.76 3.21 7.01

the fallen trees there is a rapid colonization of lianas that almost completely cover herbs and shrubs.

Other small mammals captured on the same sampling erid during | or both of the same sampling sessions were Gracilinanus agilis, Monodelphis kunsi, and Marmosa paraguayana (family Didelphidae), and Akodon montensis, Cerradomys maracajuensis, Hylaeamys megacephalus, Oligoryzomys mattogrossae, O. nigripes, and Rhipidomys macrurus (subfamily Sigmodontinae). Calomys callosus, Euryoryzomys russatus, Oligoryzomys cf. flavescens, and Sooretamys angouya (Sigmodontinae) were also collected on this grid, at other times.

The subadult male Kannabateomys amblyonyx (CZPLT 486) was found dead on the side of the road by a bridge over the Arroyo Palmito, in southern Paraguay, in an area of degraded Interior Atlantic Forest. The bridge Spans a stream leading to San Benito, a shallow lake approximately 650 m away. The area is surrounded by a mosaic of small-holder farms, natural grassland, and a pine plantation, and a narrow corridor of degraded forest to the west incorporating mostly secondary growth flora

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Figure 4. Dorsal view of hind feet of 2 Juliomys pictipes. Left, TK 170166; right, TK 186360. Note reddish-orange coloration of dorsal hairs, and white ungual tufts, characteristic of this species.

and degraded gallery forest to the south, both containing bamboo thickets.

This species 1s known in Paraguay from | other speci- men, collected in 1902 in Sapucai, Paraguari Department (Thomas 1903), approximately 175 km northwest of our locality. A third specimen, reported to be from Capitan Meza, Itapua Department, collected by Adolfo Neun- tenfel (date not specified, but probably 1938 or 1939), is housed in the Museo Argentino de Ciencias Naturales «Bernardino Rivadavia» in Buenos Aires. Approximately 170 specimens from the same collector are listed from the same locality, and it is unclear whether they were collected there or collected in nearby localities, includ- ing northern Argentina, and shipped from Capitan Meza. Thus, we cannot confidently consider this to be a Para- guayan record for K. amblyonyx. Additionally, a reliable sight record was reported from Estancia Nueva Gambach (26.6379° S, 055.6643° W) on the southern tip of San Rafael National Park on 12 December 2011 (P. Smith, pers. comm. ).

The species is easily distinguished from all other rodents in Paraguay by its size (Table 1), generally dull buffy yellowish pelage, distinctly pale with grey and black flecks around the mouth, chin and chest, and the conformation of its hind foot, especially the toes (Fig. 6). The new specimen reported here is the first confirmed specimen from Paraguay in over a century. Endemic to

the Atlantic Forest of Paraguay, Brazil and Argentina, the southern bamboo rat is a bamboo specialist (Stall- ings et al. 1994), with an exclusive diet of shoots and leaves of bamboo, as well as using bamboo for arboreal travel (Kierulff et al. 1991, Olmos et al. 2009). They are known to live inside thickets of bamboo near to water- ways, including in planted or naturally occurring bamboo patches in close proximity to agriculture (Emmons et al. 2015).

Thomas (1903) described Kannabateomys amblyonyx pallidior based on the specimen from Sapucay, listing it as a paler subspecies of K. amblyonyx previously described from Sao Paulo, Brazil. The new specimen concurs with several of the features he described, including a general pelage of dull buffy yellowish, long and soft fur, and a pale, nearly white, venter. The head is darker than the body with less buffy yellowish coloring and more gray and black flecks; and around the mouth, chin, and chest it is paler to white. The vibrissae are long, dark and dense. The base of the tail is thickened and similar in color to the dorsal aspect of the body. From the mid-point to the tip of the tail the pelage becomes paler and white. The tail is furred to the tip, and the terminus has brown and black flecks, concurring with the description of the species by Emmons et al. (2015), rather than the white tip described for K. a. pallidior by Thomas (1903). The ventral surface of the limbs is pale yellow to white, with paler fur on

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Figure 5. Dorsal and ventral views of the crania of three Juliomys pictipes from Paraguay. Left, FACEN-CZ 014; center, TK 170166; right, TK 186360. See Table 1 for measurements of the 2 skulls.

the tips of the digits of all 4 feet. This is consistent with the description for both K. amblyonyx and its subspe- cies K. a. pallidior. However, caution should be used, as the descriptions were both based on adult animals, and our specimen is a subadult. Further geographical study is required to determine the validity of the subspecies (Emmons et al. 2015).

Discussion

Numerous authors have noted that the mammalian fauna of Paraguay is poorly known (e.g. Myers 2002, Lopez- Gonzalez et al. 2014, de la Sancha et al. 2017), and new species continue to be added to the known mammalian fauna of the country (e.g. D’Elia et al. 2008, de la Sancha et al. 2009, Owen et al. 2014). This report continues and reinforces that trend, and these records from the Upper

Parana Atlantic Forest biome of eastern Paraguay also demonstrate that new and important faunal information may still be encountered, even in areas considered to be well sampled. Canindeyti Department, which includes the Reserva Natural del Bosque Mbaracayu, is the most intensively sampled department in Paraguay, with over 6,200 mammalian specimens in collections in Europe, North and South America (unpublished data). Neverthe- less, this and several other recent reports illustrate that even in this heavily sampled area, the mammal fauna may yet be incompletely known, let alone the factors affect- ing community composition, population demographics, responses to land-use changes, and risk of zoonotic dis- ease maintenance and transmission.

Based on this realization, we make the following recommendations: (1) advanced training of more and better field-oriented biologists, to facilitate more collect-

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Figure 6. Lateral view of head, neck and forefeet (left), and ventral view of hind foot (right) of Kannabateomys amblyonyx, CZPLT 486.

ing and better field skills; (2) advanced training of more and better collections-oriented biologists, to facilitate better maintenance of scientific biological collections, and careful review of existing specimens; (3) better train- ing for administrators of permit-granting entities within governments, to facilitate (rather than obstruct) scientific collection and other fieldwork; and (4) better education of administrators of both publicly and privately administered protected areas, to engender their support for scientific research in those protected areas, especially as those become the few remaining natural laboratories available.

Acknowledgements

We thank Pablo Teta and Julio Torres for pointing out the previously collected specimen of De/omys among a group of Hylaeamys specimens. Andrea Weiler approved the loan of Juliomys pictipes specimen FACEN-CZ 014 housed at the Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Asuncion. Ana Rivarola pro- vided much of the vegetation information cited here. Numerous young biologists assisted with fieldwork, especially Julieta Sanchez, Belén Barreto, and Erica Rios. Danilo Salas, Sixto Fernandez, Arnaldo Alfonzo, and Fredy Ramirez (Fundacion Moises Bertoni, Para-

guay) were consistently supportive of our work at the RNBM. Emiliano Galeano was indispensable in our fieldwork at the RNBM. Hugo Cabral created the map for Figure 1, and Cristian Palacios did the photography and arrangements for Figures 2—5. Paul Smith kindly reviewed an earlier version of the manuscript, and pro- vided helpful suggestions, as well as sharing his sight record of K. amblyonyx and providing additional infor- mation about the K. amblyonyx collecting locality. We thank W. Corréa-Tavares and an anonymous reviewer for their comments which helped to clarify and improve this manuscript. RDO and CBJ acknowledge the support of the National Institutes of Health (NIH, U.S.A.) grant ROI AI103053. RDO was partially supported by the Programa Nacional de Incentivo a los Investigadores (CONACYT, Paraguay).

Authors’ Contributions

All authors participated in collecting 1 or more of the specimens reported here. R. Owen drafted the manuscript and took measurements on the D. dorsalis and J. pictipes specimens. L. McMahon took measurements on the K. amblyonyx specimen. All authors reviewed and assisted in revising the manuscript.

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