Research Article |
Corresponding author: Ruth Salas ( rsalas@amnh.org ) Academic editor: Alfred Wheeler
© 2018 Ruth Salas, Randall T. Schuh.
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:
Salas R, Schuh RT (2018) Macrotylus henryi, a new species of Pelargonium-feeding Cremnorrhinina from South Africa (Hemiptera, Miridae, Phylinae, Cremnorrhinini). In: Wheeler Jr AG (Ed.) A Festschrift Recognizing Thomas J. Henry for a Lifetime of Contributions to Heteropteran Systematics. ZooKeys 796: 267-280. https://doi.org/10.3897/zookeys.796.21429
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Macrotylus henryi is described as a new species from South Africa. This new taxon is recorded as feeding on species of Pelargonium (Geraniaceae) in the Western Cape. Documentation is provided in the form of diagnosis, description, habitus photographs, scanning electron micrographs, illustrations and images of genitalic structures, detailed distributional data, host plant information, and images of hosts and habitats. Morphological traits are similar to species of Macrotylus Fieber from the Northern Hemisphere, but coloration is substantially variable, and the structure of the male genitalia is distinctive.
Geraniaceae , Heteroptera , host plant, long free pulvilli, plant bug, Western Cape
Macrotylus Fieber, 1858 is now placed in Cremnorrhinini: Cremnorrhinina based on phylogenetic analysis that includes morphology and DNA sequence data (
Collecting of Miridae in South Africa produced new specimens of Macrotylus as well as host and localities documentation. Fieldwork focused on Namaqualand, the Little Karoo, and the fynbos vegetation of the Western Cape, because of the extreme botanical diversity of the area and the very limited sampling of Miridae for the area during the eight-month expedition of J.A. Slater and colleagues during 1967–1968, which had been the only concerted effort to collect phytophagous Heteroptera in South Africa.
In the present paper a new species, Macrotylus henryi, is described which feeds on plant species in the family Geraniaceae endemic to South Africa. This new taxon is dedicated to Thomas J. Henry in recognition of his contributions to our knowledge and understanding of true bug taxonomy and host associations.
Unique matrix code labels were affixed to each of 360 specimens examined; these codes are therefore referred to as “unique specimen identifiers” (USIs). The USI codes are composed of an institution and project code (AMNH_PBI) and a unique number (00393079). The AMNH_PBI prefix was removed from the Specimens Examined sections of the paper to save space and make these sections more readable, but was retained for the holotype and figures. Data for these specimens were captured using the American Museum of Natural History instance of the Arthropod Easy Capture specimen database, formerly known as Planetary Biodiversity Inventories database. Specimen data can be viewed on line through research.amnh.org/pbi/heteropteraspeciespage/ and discoverlife.org, and through the iDigBio web portal (idigbio.org/portal).
Color digital habitus images of the bugs were prepared using a Microptics-USA/Visionary Digital photomicrographic system as developed by Roy Larimer; multiple layers were stacked using Helicon Focus. Habitus photos are proportional to the size of the actual specimens so that relative sizes can be deduced from comparison of the specimen images.
Details on specimen measurements are provided in Table
Host samples were collected and pressed in the field in conjunction with each collecting event. These specimens were subsequently identified by botanical specialists; the botanical names were then associated with the individual bug specimens through a specimen database and during the labeling process. Macrotylus specimens were collected on plants in flower. Thus, the identification of the host plants is to the level of species. Host field photos were made using a Nikon D1 SLR digital camera.
Scanning electron micrographs were prepared using a Hitachi S-4700 digital SEM. Male genitalic illustrations were prepared as pencil drawings using a Nikon Eclipse 80i compound microscope, then scanned and rendered as graphics using Adobe Illustrator. All such illustrations were drawn with a 20× or 40× objective lens. Female genitalic images were taken with a 10× or 20× objective lens using a Nikon E800 compound microscope, photomicrographic attachment, and software.
The insect specimens examined in this study were provided by the following institutions, or material is deposited in them; institutional abbreviations used in the specimens examined sections and names of individuals who assisted handling the specimens are also listed:
Placed in Macrotylus Fieber based on the following characteristics: The projecting clypeus, the sparsely distributed, reclining dark common setae on the dorsum (Figs
Scanning micrographs of Macrotylus henryi (AMNH_PBI 00393017). A Lateral view of head and pronotum B Detail of pronotal setae C Vestiture on hemelytron D Thoracic pleuron, showing metathoracic spiracle (mttsp), metathoracic scent-gland auricle (sgaur), and scent-gland evaporatory area (sgev) E Lateral view of pygophore, left paramere (lp), phallotheca (phl), and right paramere (rp) F Lateral view of pretarsus (pul, pulvillus).
Distinguished from other South African Macrotylus species by its larger size, vestiture type, and the structure of endosoma. Macrotylus niger mostly black; M. hemizygiae, although often yellow green as in some M. henryi specimens, with shining woolly setae as well as densely placed dark setae on dorsum. These two previously described South African species apparently lacking denticles seen on apex of endosoma in M. henryi (Fig.
Male. Relatively large, total length 4.85–6.26, pronotum width 1.20–1.56 (Table
Surface and vestiture (Figs
Structure (Figs
Thorax: Pronotum wider than long, slightly campanulate, posterior lobe weakly elevated; thoracic pleuron with sericeous setae and scattered common setae, metathoracic scent-gland evaporatory area triangular (Fig.
Genitalia: Pygophore (Fig.
Female. Coloration, surface, and vestiture (Fig.
Macrotylus henryi, digital female genitalic images (AMNH_PBI 00393199) showing the bursa copulatrix and the bursa copulatix. Abbreviations: acgl, accessory gland; bsfg, basal sclerite of first gonapophyses; dlp, dorsal labiate plate; irs, interramal sclerites; latov, lateral oviduct; mdext, medioventral extension of ventral labiate plate; mdscl, medial interramal sclerite; pw, posterior wall; sclr, sclerotized ring; vlp, ventral labiate plate.
Length | Width | Ratio | ||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Body | CunClyp | Head | Prono | Scut | Cun | AntSeg2 | Head | Prono | Scut | IntOcDi | WH/LH | WH/WP | IOD/WH | AS2/WH | WP/LP | |
♂ (N=23) Mean | 5.49 | 3.89 | 0.57 | 0.60 | 0.52 | 0.96 | 1.67 | 0.80 | 1.37 | 0.71 | 0.35 | 1.43 | 0.59 | 0.44 | 2.09 | 2.28 |
SD | 0.46 | 0.41 | 0.09 | 0.06 | 0.05 | 0.07 | 0.14 | 0.04 | 0.12 | 0.07 | 0.02 | 0.17 | 0.03 | 0.02 | 0.07 | 0.12 |
Range | 1.41 | 1.13 | 0.30 | 0.20 | 0.16 | 0.26 | 0.46 | 0.15 | 0.36 | 0.21 | 0.06 | 0.67 | 0.09 | 0.09 | 0.31 | 0.50 |
Min | 4.85 | 3.43 | 0.43 | 0.51 | 0.46 | 0.82 | 1.46 | 0.74 | 1.20 | 0.63 | 0.32 | 1.15 | 0.54 | 0.39 | 1.93 | 2.04 |
Max | 6.26 | 4.56 | 0.73 | 0.71 | 0.61 | 1.08 | 1.92 | 0.88 | 1.56 | 0.84 | 0.39 | 1.82 | 0.63 | 0.48 | 2.24 | 2.54 |
♀ (N=23) Mean | 5.47 | 3.96 | 0.59 | 0.63 | 0.53 | 0.89 | 1.62 | 0.79 | 1.45 | 0.74 | 0.41 | 1.35 | 0.54 | 0.52 | 2.06 | 2.32 |
SD | 0.48 | 0.37 | 0.07 | 0.05 | 0.05 | 0.08 | 0.11 | 0.05 | 0.11 | 0.06 | 0.02 | 0.12 | 0.02 | 0.02 | 0.06 | 0.09 |
Range | 1.45 | 1.13 | 0.28 | 0.18 | 0.17 | 0.25 | 0.43 | 0.15 | 0.33 | 0.19 | 0.06 | 0.48 | 0.06 | 0.07 | 0.27 | 0.32 |
Min | 4.74 | 3.44 | 0.45 | 0.54 | 0.45 | 0.76 | 1.37 | 0.73 | 1.30 | 0.66 | 0.38 | 1.17 | 0.51 | 0.48 | 1.88 | 2.21 |
Max | 6.19 | 4.57 | 0.72 | 0.72 | 0.62 | 1.01 | 1.80 | 0.88 | 1.63 | 0.85 | 0.44 | 1.65 | 0.57 | 0.55 | 2.16 | 2.52 |
Named for Thomas J. Henry.
Recorded from species of Pelargonium L’Hér. (Geraniaceae) (Fig.
Western Cape, from near Clanwilliam to south Cape Peninsula, and east to near Mossel Bay; from sea level to ~650 m elevation (Fig.
Digital photographs of living and pressed Pelargonium spp that are hosts of Macrotylus henryi, from Western Cape, South Africa A–CPelargonium cucullatum × betulinum (3.2 km E of Hermanus) D Randall T. Schuh collecting on P. cucullatum × betulinum (Koeel Bay, 20 km S of Strand on R44) E–G Pelargonium radens (10.5 km E of Clanwilliam, Cedarberg Range) H Christiane Weirauch collecting (3.2 km E of Hermanus).
Holotype: SOUTH AFRICA: Western Cape: 3.2 km E of Hermanus, 34.40536S, 19.32737E, 33 m, 10 Nov 2003, Schuh, Cassis, Weirauch, Pelargonium cucullatum × betulinum (L.) L’Hér. ex Aiton (Geraniaceae), det. K. Roux NYBG VOUCHER, 1♂ (AMNH_PBI 00393026) (
Paratypes: SOUTH AFRICA: Western Cape: 3.2 km E of Hermanus, 34.40536S, 19.32737E, 33 m, 10 Nov 2003, Schuh, Cassis, Weirauch, Pelargonium cucullatum × betulinum (L.) L’Hér. ex Aiton (Geraniaceae), det. K. Roux NYBG VOUCHER, 16♂ (00388606–00388621), 28♀ (00388622–00388649) (
SOUTH AFRICA: Western Cape: 3.2 km E of Hermanus, 34.40536S, 19.32737E, 33 m, 10 Nov 2003, Schuh, Cassis, Weirauch, Pelargonium cucullatum × betulinum L’Hér. ex Aiton (Geraniaceae), det. K. Roux NYBG VOUCHER, 2 nymphs (00393032, 00393033) (
The structure of the male genitalia shows essentially no variation across the range of specimens we include in M. henryi; we therefore treat all specimens under a single species in spite of the substantial variation in color and size. The endosoma is similar to that seen in many species of Macrotylus from the Northern Hemisphere, as well as the Cremnorrhinina more broadly, including particularly the genera Halophylus Schuh and Schwartz and Pulvillophylus Schuh and Schwartz from Australia.
Among members of the South African Phylinae, the habit of feeding on Geraniaceae is not shared with any other species, although species of Dicyphini are frequently encountered on that plant group.
Most species of Macrotylus from the Northern Hemisphere have been reported to feed on Lamiaceae, with some taxa on Rosaceae and Asteraceae (
In the Balkan Peninsula, Cremnorrhinus basalis Reuter is strictly associated with ephemeral Geranium rotundifolium L. (
Macrotylus henryi seems to be restricted geographically to the Western Cape (Fig.
The distribution of M. henryi is similar to that seen in the well-collected Pseudosthenarus ater Poppius and P. brendae Schuh and Salas (
With regard to the distribution of Macrotylus more broadly, there appears to be a broad disjunction on the African continent between the larger and much better known Palearctic fauna and the Macrotylus species from southern Africa.
Host plants and localities of Macrotylus henryi in Western Cape, South Africa.
Host taxon | Locality | Insect specs. |
---|---|---|
GERANIACEAE | ||
Pelargonium alternans | Western Cape: ca. 5 km E of de Doorns S of N1 | 4 |
Pelargonium cucullatum × betulinum | Western Cape: 3.2 km E of Hermanus | 129 |
Western Cape: Koeel Bay, 20 km S of Strand on R44 | 1 | |
Pelargonium cucullatum subsp. tabulare | Western Cape: Table Mountain National Park, Cape Peninsula | 62 |
Western Cape: Table Mountain National Park, Cape Peninsula, Circular Drive | 14 | |
Pelargonium radens | Western Cape: ca. 25 km E of Clanwilliam, on plains below Pakhuispas | 60 |
Western Cape: 10.5 km E of Clanwilliam, Cedarberg Range | 18 | |
Pelargonium scabrum | Western Cape: 5.6 km W of Clanwilliam on Rt 364 to Lambert’s Bay | 45 |
Western Cape: Nieuwoudts Pass, ~30 km N of Citrusdal on road to Algeria | 23 | |
Western Cape: 57.5 km NW of N2 on R327 beyond Herbertsdale | 3 | |
Unknown | Western Cape: Farm Dwars rivier, Cedarberg | 1 |
TOTAL | 360 |
Fieldwork for this project was supported by a National Science Foundation (NSF) Planetary Biodiversity Inventories award (DEB-0316495) to Randall T. Schuh and Gerasimos Cassis for the study of the Miridae subfamilies Orthotylinae and Phylinae. We thank Timothy Crowe, Chris Tobler, and Mike Picker of the University of Cape Town, Gerhard Prinsloo, Plant Protection Research Institute, Pretoria, and Lorenzo Prendini, American Museum of Natural History for logistical assistance in the conduct of field work in South Africa. We also thank the relevant authorities in the Western Cape Province for the issuance of collecting permits. Host plants were identified by the staff at the Compton Herbarium, Kirstenbosch Gardens, Cape Town, South Africa. We especially thank Edwina Marinus for her assistance in coordinating the acquisition of these data, and Koos Roux for the actual identifications. We thank Stephen Thurston, American Museum of Natural History, for assembling the figures in digital form for publication; he also prepared the digital rendering of the male genitalic illustrations and some of the habitus photos. Michael D. Schwartz, Canadian National Insect Collection, dissected the female genitalia and made the digital images. Many thanks to the editor Alfred G. Wheeler Jr., Clemson University, and reviewer Fedor V. Konstantinov, St. Petersburg State University, for their comments and suggestions on the manuscript.