Biodiversity Data Journal :
Taxonomic paper
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Corresponding author:
Academic editor: Pierfilippo Cerretti
Received: 26 Jan 2015 | Accepted: 20 Mar 2015 | Published: 25 Mar 2015
© 2015 AJ Fleming, D. Monty Wood, Daniel Janzen, Winnie Hallwachs, M. Alex Smith
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.
Citation:
Fleming A, Wood D, Janzen D, Hallwachs W, Smith MA (2015) Seven new species of Spathidexia Townsend (Diptera: Tachinidae) reared from caterpillars in Area de Conservación Guanacaste, Costa Rica. Biodiversity Data Journal 3: e4597. https://doi.org/10.3897/BDJ.3.e4597
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We describe seven new species of Spathidexia (Diptera: Tachinidae) reared from Area de Conservación Guanacaste (ACG), northwestern Costa Rica. All were reared from various species of ACG caterpillars during an ongoing inventory of caterpillars, their food plants and their parasitoids. By coupling morphology, photographic documentation, life history and molecular data, we provide a clear and concise description of each species. All are known to be previously undescribed as a result of a comprehensive study of the genus by DMW. Spathidexia atripalpus sp. n., Spathidexia juanvialesi sp. n., Spathidexia marioburgosi sp. n., Spathidexia luisrobertogallegosi sp. n., Spathidexia luteola sp. n., Spathidexia hernanrodriguezi sp. n. and Spathidexia aurantiaca sp. n. are all authored and described by Fleming and Wood. Minthodexiopsis Townsend is proposed by Wood as a new synonym of Spathidexia. A new combination proposed by Wood as a result of the new synonymy is S. flavicornis (Brauer & Bergenstamm) comb. n.
Spathidexia, Diptera, Tachinidae, tropical rain forest, tropical dry forest, parasitoid fly, host-specificity, caterpillars.
The tachinid genus Spathidexia
This work aims to build on the existing knowledge and describes seven new species of Spathidexia, all reared from caterpillars collected in ACG. In fact, all species of Spathidexia reared in ACG have been found to be undescribed species. These species are recognized based on differences in external morphology, and COI (coxI or cytochrome oxidase I) gene sequences (a.k.a. “DNA barcodes”). By coupling COI data with morphological descriptions we are able to show that abdominal markings are not only different between males and females but are also consistent within a species. As such, they are ideal tools for visual species differentiation.
The treatments reported here is focused on placing names on the species reared from ACG, thereby preparing them for later detailed ecological and behavioral accounts and studies that will normally extend across ACG ecological groups, whole ecosystems, and taxonomic assemblages much larger than a genus. However, all ACG Spathidexia reared to date in ACG are parasitoids of monocot-eating caterpillars in the Nymphalidae (Satyrinae) or Hesperiidae (Hesperiinae).
BMNH The Natural History Museum, London, United Kingdom
CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada
USNM National Museum of Natural History, Washington, D.C., USA
INBio Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica
NHMW Naturhistorisches Museum, Wien, Austria
MZUT Musei di Zoologia, Università di Torino, Torino, Italia
CAS California Academy of Sciences, San Francisco, CA, USA
All flies and rearing information described here were found by the 35+ year-old ongoing inventory of the caterpillars, their food plants and their parasitoids of the dry forest, rain forest, cloud forest, and intergrades, in the 125,000+ ha terrestrial portion of Area de Conservación Guanacaste (ACG) in northwestern Costa Rica (
In brief, caterpillars at all instars (and sometimes pupae) are found in the wild by a wide variety of search methods, and reared in captivity on the food plant species on which they were found, until they produce an adult, a parasitoid, or die of other causes. Each caterpillar is documented as an individual, as are the adult parasitic flies.
This inventory has reared about 600,000 wild-caught caterpillars since 1978. All frequencies of parasitism reported here need to be considered against this background inventory. Equally, it is patently obvious that the inventory searches some kinds of vegetation and height off the ground much more thoroughly than others, and it also searches throughout the year. Comparison of reared species of parasitoids with those collected by net or Malaise traps demonstrates that, to date, the caterpillar inventory has encountered well less than half the species of caterpillar parasitoids present in ACG. The largest unsampled void is the upper foliage of the canopy above about 3–4 m above the ground.
Our descriptions of new species are deliberately brief and only include some differentiating descriptions of body parts and colors that are commonly used in tachinid identification. These brief descriptions are complemented with an extensive series of color photos of every species to illustrate the readily-observed differences among these species.
Habitus photographs were taken using a Canon T3i digital SLR, using a 65mm Macro Photo Lens 1:2.8 (MP–E 65mm), mounted on a microscope track stand (AmScope, Model: TS200) modified to accept a Manfrotto QR 200PL–14 quick release plate. Images were shot in aperture priority, allowing the camera to control shutter speed at f/4.5. Stacks of 40 images were taken at equal distance increments. Illumination was provided with a homemade reflective dome (instruction for dome creation can be found at: http://www.cdfa.ca.gov/plant/ppd/entomology/Dome/kd–200.html) placed over a 144 LED ringlight (AmScope, Model: LED–144–YK).
The photographic series were processed using Photoshop CS6, and digitally stacked with Zerene Stacker Software v1.04. This produced a final composite image maximizing image quality and depth of field.
The terminology used for components of the terminalia (which refers here only to the sclerotized parts of the genitalia, and not to the soft internal structures) and other body parts, follows
All specimens listed as examined are considered paratypes, except for the holotype which is noted separately.
Wherever a specimen label has been cited, the information is presented using the following symbols: /, indicates the end of a line; //, indicates the end of a label. Labels are presented from top (closest to the specimen) to bottom, with any comments about the label being given in square brackets.
All caterpillars reared from the ACG efforts receive a unique voucher code in the format of yy–SRNP–xxxxx. Any parasitoid emerging from this caterpillar receives the same voucher code, and then if/when later the parasitoid is dealt with individually, it receives a second voucher code unique to it, in the format of DHJPARxxxxxxx. The voucher codes and collateral data assigned to both host and emergent parasitoids are available at http://janzen.bio.upenn.edu/caterpillars/database.lasso. To date, all DHJPARxxxxxxx coded tachinids have had one leg removed for attempted DNA barcoding at the Biodiversity Institute of Ontario (BIO) at the University of Guelph, with all collateral data and all successful barcodes permanently and publically deposited in the Barcode of Life Data System (BOLD, www.boldsystems.org) (
Inventoried Tachinidae were collected under Costa Rican government research permits issued to DHJ since 1978, and likewise exported under permit by DHJ from Costa Rica to Philadelphia, and then to the final depository in the Canadian National Insect collection in Ottawa, Canada. Tachinid identifications for the inventory were done by DHJ in coordination with a) visual inspection by AJF and DMW, b) DNA barcoding by BIO, MAS, and BOLD, and c) correlation with host caterpillar identifications by DHJ and WH through the inventory itself. Dates of capture of each reared fly in the inventory are the dates of eclosion of the fly, and not the date of capture of the caterpillar. This is because the fly eclosion date is much more representative of the time when that fly species is on the wing than is the time of capture of the caterpillar or (rarely) finding a parasitized pupa. However, the collector listed is the parataxonomist who found the caterpillar, rather than the person who retrieved the newly eclosed fly from its rearing bag or bottle, and processed it by freezing, pinning, labeling and oven–drying. Fly biology and degrees of parasitization by these flies will be the detailed subject of later papers.
DNA barcodes (standardised 5’ region of the mitochondrial cytochrome c oxidase I (COI) gene) for all ACG inventory specimens were obtained using DNA extracts prepared from single legs using a glass fibre protocol (
Spathidexia
Gymnopalpus
Minthodexiopsis
Minthohoughia
Stenaulacodoria
antillensis
brasiliensis
cerussata
cinereicollis
clemonsi
creolensis
cylindrica
dicta
dunningii
rasilis
elegans
flavicornis
nexa
niveomarginata
pallida
reinhardi
setipennis
spatulata
*The new combination is proposed by DMW and result from new generic synonymy and the examination of the type material of other nominal species. Type label information is included for those species.
Male (Fig.
Female (Fig.
The presence of black palpi is its strongest diagnostic character, differentiating it from almost all other species. This species differs from S. atypica, which also has black(ish) palpi, in having 3 postsutural dorsocentral bristles, bare eyes, and the presence of small black hairs covering entire body especially prominent extending from anepimeron and underside of T1+2. Differs, from S. antillensis, which also bears two bold thoracic vittae but does not bear black palpi.
From the Latin “ater”, color black, and the noun “palpus”, literally the palm of the hand, in reference to the black palpi present in both sexes of this species.
Costa Rica, ACG, Prov. Guanacaste, rain forest and rain forest/dry forest ecotone, 290-580m elevation.
Reared from four species of grass-eating rain forest satyrine Nymphalidae (4 records). One larva per host.
Male (Fig.
Female (Fig.
Differs from S. cerussata, who also possesses fine white hairs on parafacialia, due to its broadly yellow side coloration, and yellow orange antenna.
This species is named in honor of Sr. Mario Burgos Cespedes (RIP), whose large land holdings in Sector Pocosol became major parts of the newly forming Area de Conservacion Guanacaste in the late 1980's.
Costa Rica, ACG, Prov. Alajuela and Guanacaste, rain forest and dry forest, 96–1140 m elevation.
Reared 221 times from 5 species of Talides (Hesperiidae; Hesperiinae), all feeding on broad-leafed rain forest monocots (Heliconia spp., Heliconiaceae and Musa spp., introduced Musaceae). Usually 5–12 larvae per caterpillar. This frequently-reared species of Spathidexia displays a very shallow DNA barcode split into two groups (see Supplemental Appendix 1) that are identical in morphology, microgeographic location in ACG, and host records. We judge these specimens to be one species of Spathidexia but they may merit a subsequent deeper genetic probe.
Male (Fig.
Female (Fig.
Differs from S. cerussata, who also possesses fine white hairs on parafacialia, due to its gold thorax, broadly yellow lateral coloration, and yellow orange antenna with darkened apices.
This species is named in honor of Sr. Luis Roberto Gallegos (RIP), whose large land holdings in Sector El Hacha, Sector Orosi and Sector Pocosol became major parts of the newly forming Area de Conservacion Guanacaste in the late 1980's.
Costa Rica, ACG, Prov. Guanacaste, rain forest, cloud forest (marginally) and dry forest-rain forest interface (primarily), 280–1460 m elevation.
Reared 50 times from Conga chydea (Hesperiidae, Hesperiinae) only, always feeding on grasses on forest edges in rain forest and the interface between rain forest and dry forest.
Male (Fig.
Female (Fig.
Species closely resembling S. clemonsi, differs from the former due to the presence of yellow tibiae on the front pair of legs, yellow/pale hairs covering the entire body, and costal spine is reduced in S. luteola, not absent as in S. clemonsi. This species is quite similar to Spathidexia brasiliensis, from which it can be differentiated by the former having an orange pedicel and narrow ocellar triangle.
From the Latin adjective, “luteus”, yellow, in reference to the pale yellow hairs covering the body of this species, particularly prominent on the underside of the abdomen.
Costa Rica, ACG, Prov. Guanacaste, dry forest and dry forest-rain forest interface, 300–600 m elevation.
Reared from 2 species of Hesperiidae, Hesperiinae, eating Lasiacis (Poaceae) (2 records), bearing 1–5 larvae per caterpillar.
Male (Fig.
Female (Fig.
Resembles S. cerussata, differs from the former in the presence of apically enlarged palpus, thorax with almost indistinguisheable vittae, a haired wing vein R1, and orange pedicel. Females in S. cerussata are similarly colored to males a characteristic not shared by S. hernanrodriguezi.
This species is named in honor of Sr. Hernan Rodriguez Arce (RIP), whose Finca San Gerardo became a large part of Sector San Cristobal of the newly forming Area de Conservación Guanacaste in the early 1990's.
Costa Rica, ACG, Guanacaste, dry forest, 155–540 m elevation.
Reared from 3 species of Hesperiidae, Hesperiinae (4 records), feeding on grasses. 5–12 larvae per caterpillar.
Male (Fig.
Female (Fig.
This species differs from the only other reddish-orange species in Spathidexia, S. creolensis from the southern US in that Spathidexia aurantiaca has more pronounced pruinosity along abdominal margins, black ground color on T5 in both sexes, and gold along the fronto-orbital plate.
From Latin, "aurantiaco" referring to the orange ground color of the abdomen.
Costa Rica, ACG, Prov. Alajuela, rain forest, 340–420 m elevation.
Reared from Hesperiidae, specifically two species of sedge (Cyperaceae)-eating Hesperiinae (4 records). 1–5 larvae per caterpillar.
Female(Fig.
Similar to S. setipennis in the thoracic markings, however S. juanvialesi is strongly different in the coloration of the antenna, and the extent of the yellow on the legs.
This species is named in honor of Sr. Juan Viales (RIP), whose Finca San Cristobal became a large part of Sector San Cristobal of the newly forming Area de Conservación Guanacaste in the eary 1990's.
Costa Rica, ACG, Prov. Guanacaste, rain forest, 440–580 m elevation.
Reared twice from grass-eating rain forest Hesperiinae (Hesperiidae).
Key to the species of Spathidexia reared from Area de Conservación Guanacaste, Northwestern Costa Rica This key was prepared based on the specimens collected as a result of the 40+ year rearing effort conducted in ACG. Our key is intended to speak to the fauna present within the confines of the ACG and is not meant to be an exhaustive key to the species within the confines of Costa Rica. |
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1 | Proclinate orbital bristles absent | 2 |
– | Proclinate orbital bristles present | 4 |
2 | Fronto-orbital plate entirely silver (no trace of gold)(Figs |
3 |
– | Fronto-orbital plate bearing at least traces of gold at least at apex, and surrounding ocellar triangle (Figs |
Spathidexia luteola sp. n. (♂) |
3 | Thoracic vittae black, prominent; vein R1 bare (Fig. |
Spathidexia atripalpus sp. n. (♂) |
– | Grey thorax with greyish indistinct vittae (only slightly visible, Fig. |
Spathidexia marioburgosi sp. n. (♂) |
4 | Ground color of abdomen dark, but bearing lateral orange/yellow blotches; or mostly orange (Figs |
5 |
– | Abdominal tergites black, with grayish tomentose bands covering up to basal 1/3 of tergal surface, and with no traces of orange/yellow blotching on abdomen (Figs |
9 |
5 | Ground color of abdomen dark, but bearing lateral orange/yellow blotches (Figs |
6 |
– | Ground color of abdomen orange along its entirety; or with darkened T5 in the case of females (Fig. |
Spathidexia aurantiaca sp. n. (♂ & ♀) |
6 | Abdomen bearing orange/yellow blotches along sides of abdominal tergites, (T1+2, T3, T4, and sometimes T5) (Figs |
7 |
– | Abdomen with yellow blotches only on T1+2 (Fig. |
Spathidexia atripalpus sp. n. (♀) |
7 | Fronto-orbital plate entirely silver with no trace of gold, or if gold present, then only in trace amount alongside ocellar triangle (Figs |
8 |
– | Fronto-orbital plate bearing traces of gold at least at apex, and surrounding ocellar triangle up to insertion of first proclinate orbital bristle (Fig. |
Spathidexia luteola sp. n. (♀) |
8 | Fronto-orbital plate entirely silver with no trace of gold; wide light brown frontal vitta; antenna black only orange on pedicel (Fig. |
Spathidexia juanvialesi sp. n. (♀) |
– | Fronto-orbital plate entirely silver with slight trace of gold alongside ocellar triangle; narrow black frontal vitta; antenna bright orange (Fig. |
Spathidexia marioburgosi sp. n. (♀) |
9 | Fronto-orbital plate with gold color only at apex surrounding ocellar triangle, descending to insertion of upper proclinate orbital in the case of females; antenna dark, only pedicel appears orange (Fig. |
Spathidexia hernanrodriguezi sp. n. (♂ & ♀) |
– | Fronto-orbital plate with gold color extending up to second proclinate orbital bristle; antenna orange (Fig. |
Spathidexia luisgallegosi sp. n. (♂ & ♀) |
We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (Janzen et al. 2009, Janzen & Hallwachs 2011) who found and reared the specimens used in this study, and the team of biodiversity managers who protect and manage the ACG forests that host these tachinids and their caterpillar hosts. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699, and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, Permian Global, and University of Pennsylvania (DHJ & WH). This study has been supported by the Government of Canada through its ongoing support of the Canadian National Collection, Genome Canada, the Biodiversity Institute of Ontario, and the Ontario Genomics Institute (2008-0GI-ICI-03) (MAS), and by a Discovery Grant from Natural Sciences and Engineering Research Council of Canada (MAS). The authors also wish to acknowledge the efforts and time of the editor and reviewers, who provided insights and suggestions allowing us to make our work better.